Information

Are polyploidal organisms parasatized by selfish genes? And if not, why not?


Certain prokaryotes are polyploids. For example, Thermus thermophilus has about five copies of its genome (http://jb.asm.org/content/192/20/5499.full). One extreme polyploid, Epulopiscium fishelsoni, has tens of thousands of copies of its genome (http://www.pnas.org/content/105/18/6730.full). (Some eukaryotes are polyploid as well.)

As each copy replicates separately it would seem likely that spontaneous mutations would arise and over time, and such an organism would become a hybrid of many distinct genomes coexisting in a single cell. This seems like an ideal place for selfish copies to arise (though perhaps no more so than any other organism - I think I've read that even prokaryotes are prone to plasmid parasites).

So, does this happen? If not, why not? Is there some copy-checking mechanism (some extremeophiles seem to have a DNA-repair mechanism) or some mechanism that inhibits reproduction by genomes that are (somehow) determined to not be contributing to the overall metabolism of the cell?


In short, being haploid or polyploid should not protect you from Selfish Genetic Elements (SGES).

Almost all eukaryotes are polyploid. Plants are the oft-cited example. Wheat, I believe, is hexaploid for example. Polyploid bacteria are the exceptions typically (if we consider ploidy to be of the whole genome). However, even in truly haploid bacteria, the genome in any given cell is usually actively being replicated, so some of the genes are actually present more than once.

Regardless, replication and repair is an enormously well controlled and regulated process: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1479556/


SGEs can take many forms depending on your definition and are ubiquitous in nature as far as we can tell. Plasmids which harbour addiction factors, insertion elements, transposons, prophages etc are all technically SGEs. See here for a review: https://www.ncbi.nlm.nih.gov/pubmed/21227262

I can't find any literature on SGEs in polyploidy directly, but it is well documented that bacteria harbour a considerable amount of selfish DNA - it seems likely that this will be the same if not worse in polyploidy. Plasmid detection is a difficult computational and experimental process (http://bioinformatics.oxfordjournals.org/content/early/2016/11/18/bioinformatics.btw651.full) but one of the hallmarks of plasmid sequences are typically addiction factors (a gene that the host cell must maintain in order to survive - usually an anti-toxin to neutralise a toxin present on a different DNA molecule) as well as phage genes.


To address a couple of things specifically:

As each copy replicates separately it would seem likely that spontaneous mutations would arise and over time

This is somewhat true, there is always minor variation in a population (else evolution wouldnt work!). Sometimes you'll find this is also the case in a single cell - but less often. All cells we have found so far have DNA replication and repair mechanisms - https://en.wikipedia.org/wiki/DNA_repair. The frequency of mutation in the genome is not directly linked to the amount of selfish elements though, making your second point contentious:

This seems like an ideal place for selfish copies to arise

I agree with the premise of polyploidy probably inviting more SGEs (thinking specifically of IS elements and transposons here really), but not because of mutation - rather, just because there are more DNA molecules and sites present for those elements to incorporate into.


Polyploidy in tissue homeostasis and regeneration

Polyploid cells, which contain multiple copies of the typically diploid genome, are widespread in plants and animals. Polyploidization can be developmentally programmed or stress induced, and arises from either cell-cell fusion or a process known as endoreplication, in which cells replicate their DNA but either fail to complete cytokinesis or to progress through M phase entirely. Polyploidization offers cells several potential fitness benefits, including the ability to increase cell size and biomass production without disrupting cell and tissue structure, and allowing improved cell longevity through higher tolerance to genomic stress and apoptotic signals. Accordingly, recent studies have uncovered crucial roles for polyploidization in compensatory cell growth during tissue regeneration in the heart, liver, epidermis and intestine. Here, we review current knowledge of the molecular pathways that generate polyploidy and discuss how polyploidization is used in tissue repair and regeneration.


General Overview

Although interest in phenotypic plasticity has increased since the 1980s (Forsman 2014), the resulting proliferation of the literature can make it difficult to keep concepts organized. As a starting point, Whitman and Agrawal 2009 should be the first piece read in this entire bibliography because of its accessibility. A common problem when sifting through literature on phenotypic plasticity is the terminology. The authors of Fusco and Minelli 2010 do a good job of defining and clarifying terms in their review. Stearns 1989 provides a historical perspective in terms of the writing, but also provides a brief history of the field (to that point). While a lot of plasticity research focuses on “higher-order” phenotypes, Callahan, et al. 1997 sets the stage for current work on phenotypic plasticity at the molecular level, which is complemented by Piersma and Drent 2003, which emphasizes how physiological processes are phenotypically plastic. Wund 2012 and Forsman 2014 are particularly useful as introductions to the types of questions being asked by plasticity researchers and how one can test hypotheses relating to plasticity. Finally, Nijhout 1990 provides an excellent introduction to the thinking that drives much of the current research being done on plasticity and evolution—particularly that development is completed by genes alone.

Callahan, H. S., M. Pigliucci, and C. D. Schlichting. 1997. Developmental phenotypic plasticity: Where ecology and evolution meet molecular biology. BioEssays 19:519–525.

This review highlights how evolutionary and molecular biology can successfully be married to investigate questions of developmental plasticity. It uses the phytochrome-mediated shade-avoidance and light-seeking responses of flowering plants as a model for this approach.

Forsman, A. 2014. Rethinking phenotypic plasticity and its consequences for individuals, populations, and species. Heredity 115:276–284.

The major strength of this review is its emphasis on how to study and test hypotheses relating to phenotypic plasticity. It provides a “whole-organism” rather than “single-trait” perspective for understanding plasticity.

Fusco, G., and A. Minelli. 2010. Phenotypic plasticity in development and evolution: Facts and concepts. In Special issue: From polyphenism to complex metazoan life cycles. Edited by G. Fusco and A. Minelli. Philosophical Transactions of the Royal Society B 365:547–556.

As an introduction to a special volume in the Philosophical Transactions of the Royal Society, this article provides one of the more succinct overviews of phenotypic plasticity, how it evolves, and its role in evolution. It is useful in trying to sort out the myriad of terminology associated with phenotypic plasticity.

Nijhout, H. F. 1990. Problems and paradigms: Metaphors and the role of genes in development. BioEssays 12:441–446.

An excellent review of how a gene-driven view of development is flawed. Although it gets a bit technical in places, this generally accessible essay should be read by anyone interested in genetics, development, and/or evolution.

Piersma, T., and J. Drent. 2003. Phenotypic flexibility and the evolution of organismal design. Trends in Ecology and Evolution 18:228–233.

By focusing on reversible forms of phenotypic plasticity (termed phenotypic flexibility), Piersma and Drent offer a perspective that highlights the ubiquity of phenotypic plasticity. They utilize less frequently noted examples of plasticity, pay particular attention to diet-induced changes in body and organ size and life-cycle staging, and emphasize the role of intra-individual variation.

Stearns, S. C. 1989. The evolutionary significance of phenotypic plasticity. BioScience 39:436–445.

As an introductory article to a special issue of BioScience, this work sets the stage for the rest of its volume, but also introduces readers to terms and concepts used throughout plasticity literature. Among other things, it gives a short, informative history of the reaction norm concept in evolutionary biology.

Whitman, D. W., and A. A. Agrawal. 2009. What is phenotypic plasticity and why is it important? In Phenotypic plasticity of insects: Mechanisms and consequences. Edited by D. W. Whitman and T. N. Ananthakrishna, 1–63. Boca Raton, FL: CRC.

This book chapter should be the first piece read by anyone interested in phenotypic plasticity. Although this chapter is in a book on insects, it provides one of the most accessible, comprehensive overviews of phenotypic plasticity available.

Wund, M. A. 2012. Assessing the impacts of phenotypic plasticity on evolution. Integrative and Comparative Biology 52:5–15.

This review (in particular Table 1) is a vital resource for those interested in testing hypotheses related to phenotypic plasticity. Almost acting as a “how-to” guide, it outlines approaches and empirical examples for testing eight key hypotheses of plasticity’s role in evolution.

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Genome Size

Extragenic DNA

The C-value paradox (like any paradox) was a product of incomplete understanding, in the case of genome structure. A major breakthrough in our understanding of genome size variation, and the solution of the C-value paradox, came with the discovery that the genomes of eukaryotes (as well as some prokaryotes) contain large and varying amounts of sequences that do not function as protein-coding genes and have no other known function other than contributing to the structure of chromosomes. This ‘extragenic’ DNA contains a wide variety of different kinds of repetitive DNA and nonrepetitive sequences. Among genomes that have been studied, two main types of repetitive sequences have been found: tandem repeats and interspersed repeats. The tandem repeats are classified according to their sedimentation profiles as satellite DNA, minisatellites, and microsatellites. Interspersed repeats include short-interspersed nuclear elements (SINEs) and long-interspersed nuclear elements (LINEs). There is growing evidence that many of these sequences represent mutated copies of active elements, including both pseudogenes (regular coding genes that have become inactive due to mutation) and mobile elements such as transposons and retrotransposons. Some of these sequences have become extraordinarily abundant in particular genomes. For example, the human genome contains about a million copies of 300 bp-long SINE retrotransposons called Alu elements, representing over 10% of the genome. Overall, extragenic DNA constitutes an astonishingly large proportion of the genome, representing at least 98% of the human genome, for example, and over 99.9% of the genome of the mudpuppy salamander (N. maculosus) ( Figure 3 )!

Figure 3 . Genic vs. ‘junk’ DNA in a salamander compared with a human.


The Selfish Gene Does Not Exist

Early in The Selfish Gene, the book’s author, Richard Dawkins, declares, “I might say to you ‘Darwin was the greatest man who has ever lived’, and you might say, ‘No, Newton was’, but I hope we would not prolong the argument.” [1] From the fact that he has chosen two scientists as the greatest human beings ever to have lived, and of the two, has ranked Darwin above Newton, it is clear that Dawkins has a very high estimation of Charles Darwin and his Theory of Evolution by natural selection.

I wonder to what extent the similarity in their last names influenced Dawkins, thus determining his future interests and beliefs, when he first heard or read Darwin’s name and his wondrous Theory of Evolution. After all, the concept of kinship, or the degree of genetic relatedness between two organisms, is fundamental to Dawkins’ selfish-gene theory. In orthographical terms, “Daw” is clearly kin to “Dar” – or phrased slightly differently, Daw’s kin to Dar-win.

When Richard was still a young boy,
he heard about Darwin the goy.
Their names almost rhymed
and thus intertwined,
he set out proclaiming, “Ahoy!

“My master, Charles Darwin, was right
when he said that most species fight.
So prepare to bear witness –
else prove your genes’ fitness,
or flee like a coward in flight!”

Besides the kinship between “Dawkins” and “Darwin,” Richard’s surname also bears a high degree of kinship with that of another famous English writer, Charles Dickens. As we shall see in the course of this essay, Richard Dawkins is the intellectual progeny of two prominent Victorian-era Charlies – Darwin and Dickens. [2] But like most unions between individuals of two distinct species, such as a horse and an ass, this intellectual union between a writer of non-fiction and another of fiction has also been sterile. Just as Dickens wrote many popular fictionalized versions of English life in the nineteenth century, Dawkins has likewise provided the world with a fictionalized version of the ultimate purpose and meaning of life: we are all here merely to serve, propagate, and perpetuate our genes, which, according to the Dawkinsian version of the biological world, are IMMORTAL.

Fundamental to Dawkins’ theory is his redefinition of the relationship between genes and the organisms they give rise to: “We were built as gene machines, created to pass on our genes.” (199) Turning the conventional view on its head, Dawkins declares that all living organisms are merely “survival vehicles” for their genes, meaning that organisms are merely the genes’ way of perpetuating themselves. Most people would object that this attribution reverses the true relationship between genes and organisms. Phrased in terms of chickens and eggs, it is like saying that the chicken is merely the egg’s way of perpetuating itself. And indeed, one way of considering Dawkins’ selfish-gene theory is by asking, “Which came first, living organisms or DNA?” [3] There is no doubt what Dawkins’ answer to this question is, since he calls DNA, or their molecular predecessors, the primordial replicators.

Dawkins begins the presentation of his novel and striking thesis by arguing that, before there was any life on Earth, there first developed molecules that were able to replicate themselves. These molecules he calls replicators, and, according to him, they were the ancestors of the DNA that determine the shapes, sizes, colours, behaviours, and all the other qualities and characteristics of all living things that have ever existed, continue to exist, and will exist in the future on the Earth. With the passage of time, according to Dawkins, some of these molecules developed a protective membrane, within which they developed and replicated themselves, as well as the organisms to which they gave rise. He declares, “the story of the replicator molecules probably happened something like the way I am telling it, regardless of whether we choose to call them ‘living’.” (18)

But how can Dawkins be so sure of himself? Was he there to witness the events that he is describing? Clearly not, since he isn’t that old and neither is he omniscient. Is there any evidence to support his account? No, for there exist no fossil remains of these microscopic replicator molecules. In other words, all that Dawkins has done is to present to his readers a plausible account of how DNA originally developed – an account, moreover, that cannot be proven because there is not, and never can be, any evidence to support it. His account connects the dots between what we do know or believe – that, in the beginning, there was no life on Earth, and then, very gradually, there developed single-celled organisms, which later developed into larger multi-celled organisms.

Since Dawkins claims to be a scientist, he should know better than to try to pawn off on his readers mere conjecture, no matter how plausible it may seem, as fact. In other words, mere plausibility has no claim to truth in science, for there are a great many things that seem plausible but are in fact wrong. [4] There are also many things that seem implausible, at least to our limited human understandings, but do in fact exist or occur in the Universe. Science has been able to establish a generally-accepted body of knowledge precisely because it has rejected mere plausibility as a sufficient guarantee of truth. Prior to its advent, unsubstantiated plausibility was taken by many people – and still is taken by many people today – to be sufficient grounds for believing that something is true.

In presenting his story of how replicator molecules became the progenitors of DNA, Richard Dawkins has propagated a pseudo-scientific creation myth that is on a par with other creation myths like the one contained in the Book of Genesis. And this is not changed by his beginning with, and basing his myth on, scientific facts that most people would accept as being true. For the truth is that, Dawkins’ hypothesis notwithstanding, we do not know how DNA first developed and gave rise to Life, just as we still do not know how and why Life first developed on the Earth, and, at least as far as we have been able to determine, nowhere else in our solar system.

Although the rejection of Dawkins’ plausible but wholly unproven – and unverifiable – account is not fatal to his selfish-gene theory, this fictional account forms an important pillar to it. For if, as he claims, genes and DNA existed before the first living creature existed, which probably was a single-celled organism, then they have a sort of regal precedence in the hierarchy of life: not only do we and all other living organisms owe all our characteristics to our genes, we would not be around if they had not first developed and began replicating themselves in the primeval, swampy, molecular soup, thus establishing an organic order in the otherwise barren, disordered, unsettled, and inorganic early conditions of the Earth.

Another aspect of the particulateness of the gene is that it does not grow senile it is no more likely to die when it is a million years old than when it is only a hundred. It leaps from body to body down the generations, manipulating body after body in its own way and for its own ends, abandoning a succession of mortal bodies before they sink in senility and death.

The genes are the immortals, or rather, they are defined as genetic entities that come close to deserving the title. We, the individual survival machines in the world, can expect to live a few more decades. But the genes in the world have an expectation of life that must be measured not in decades but in thousands and millions of years.

In sexually reproducing species, the individual is too large and too temporary a genetic unit to qualify as a significant unit of natural selection. The group of individuals is an even larger unit. Genetically speaking, individuals and groups are like clouds in the sky or dust-storms in the desert. They are temporary aggregations or federations. They are not stable through evolutionary time. Populations may last a long while, but they are constantly blending with other populations and so losing their identity. They are also subject to evolutionary change from within. A population is not a discrete enough entity to be a unit of natural selection, not stable and unitary enough to be ‘selected’ in preference to another population. (34)

Those who are familiar with Plato’s philosophy should be able to recognize what Dawkins is looking for: he is attempting to find the essence of life, the unchanging unity or identity amidst all this decay and fleeting temporality that we call Life. In other words, Richard Dawkins is searching for – and believes he has found – a biological Platonic universal that can be said to endure forever, just as Plato believed that his universals or Ideas were timeless and unchanging. The comparison is exact, for, just as Plato declared that, although all the particular manifestations of any universal, such as TREE, are temporal, since no individual tree lives forever, the universal TREE is immortal. Similarly, although Dawkins readily admits that individual genes do not live very long, a few months at the most, the GENES of which they are the particular manifestations or copies are immortal.

However, as I have argued elsewhere, specifically in the discussion of our ability to perceive similarities in The Theory of Imitation, Platonic universals do not exist. The belief in Platonic universals arose from our particular human perceptual system, whose primary function is to enable us to survive and procreate. And being able to recognize similarities and categorize objects obviously does help us to accomplish these goals. Hence, in considering any system of classification, or way of viewing certain aspects of the world, including Dawkins’ redefinition of the relationship between genes and the organisms they give rise to, the primary consideration is whether it is useful or not, and not, as Dawkins believes, whether it is true or not, since his redefinition is merely a particular way of perceiving and categorizing certain aspects of the world, among the many other possible ways of perceiving these aspects or categorizing them.

By giving such great precedence to DNA, Dawkins elevates them to a superior magisterial status which, prior to his book, they had not enjoyed. As the surprising role of genes and DNA in the development of living organisms was discovered during the twentieth century, most people attributed to them the role of faithful but inanimate blueprints that do their part in the magnificent creation and evolution of Life. “But no,” saith Richard the Rational, “thou greatly underestimatest the true and noble status of our Genes for in verity these Selfish Genes are our Masters, to whom we are eternally beholden for our forms, our intelligence, our creativity, our children, for all other living creatures – nay, for our very lives and for all that is Great, Glorious, and Grand in the world. And it is precisely for these reasons that we, their temporary servile vessels, owe our fealty, allegiance, and gratitude to these generous Masters, by seeking to perpetuate them as much and as widely as is humanly or organismically possible.” In doing so, Dawkins has revived a form of ancestor worship, but cloaked in a seemingly scientific guise: these genes, which most people previously regarded as an important but inanimate form of organic matter, are in reality alive and conscious, for they seek to replicate and perpetuate themselves, in fierce competition with other genes.

Repeatedly throughout his book, while using purposive language, Dawkins declares that we are not to take this language literally. For example, in the last excerpt, he writes, “It [the gene] leaps from body to body down the generations, manipulating body after body in its own way and for its own ends, abandoning a succession of mortal bodies before they sink in senility and death.”

If we allow ourselves the licence of talking about genes as if they had conscious aims, always reassuring ourselves that we could translate our sloppy language back into respectable terms if we wanted to, we can ask the question, what is a single selfish gene trying to do? It is trying to get more numerous in the gene pool. Basically it does this by helping to program the bodies in which it finds itself to survive and to reproduce. (88)

And yet, it is entirely possible to describe the dominance of certain genes in the world without using such misleading terminology. [5] For example, he could have written instead, “as a result of natural selection, certain genes become more numerous in the gene pool, by helping organisms to survive and reproduce, and thereby pass on their genes to succeeding generations.” Unlike Dawkins’ misleading and highly problematic language, this passage does not attribute conscious aims to inanimate objects like genes. The question we should ask ourselves is, “Why does Richard Dawkins constantly use purposive language when he discusses genes?”

In Hamlet, while watching the play [6] enacted, at Hamlet’s behest, by a group of itinerant actors, the protagonist’s mother, Queen Gertrude, declares, “The Lady doth protest too much, methinks.” What she means is that the Player Queen, despite her protestations to the courtings of her deceased husband’s brother, is actually quite willing to marry him, and soon accepts him in her marital bed. Similarly, despite Dawkins’ repeated protestations, he is clearly being disingenuous, for the fact is that he wants his readers to attribute conscious aims to genes, as he himself does, for it is central to his grandiose Theory of the Immortal Gene. [7] After all, he has called his book The Selfish Gene, rather than, for example, The Dominant Gene, or How Certain Genes Become Dominant in the World, since it can happen that some things become dominant, but without seeking consciously or deliberately to become dominant.

Dawkins makes frequent use of analogies in his book to buttress his claims. There are a number of real-world analogies that will help us to see what is fundamentally wrong with his claim that all living organisms are merely “survival vehicles” for their genes. There are many similarities between the relationship between genes and organisms and the relationship between architectural blueprints and the buildings they give rise to. Someone who was wholly ignorant of the true function of buildings might make the observation that the more buildings there are, the more blueprints there are too! If this person wanted to make a startling declaration to the world, as Richard Dawkins clearly wanted to do before he wrote his book, one would then leap to the conclusion that, ergo, the true function of buildings is to propagate, perpetuate, and increase the number of blueprints in the world! In addition, the fact that no building lasts forever does not change the fact that, while they exist, buildings serve many useful purposes, such as dwellings that protect their users and inhabitants from the weather and from other human beings and animals, and as storage facilities for various things. Similarly, the fact that no living organism is immortal does not change the fact that, during the time that it exists, its existence is meaningful both to it and to other living organisms, and not merely as a subservient “survival vehicle” for its genes.

To argue, as Dawkins does, that living creatures are merely “survival vehicles” for genes is no different from saying that buildings are merely survival vehicles for blueprints. Or, to consider some other examples, that paintings are merely the paints’ way of perpetuating and multiplying themselves, since it is a fact that the more paintings there are in the world, the greater is the number of pots or tubes of paint that culinary dishes are merely the recipes’ way of perpetuating and multiplying themselves that musical compositions are merely the notes’ way of perpetuating and reproducing themselves or that books are merely survival vehicles for letters, since books contain numerous combinations of letters, and the more books there are, then obviously the greater the number of times each letter is able to reproduce, or make copies of, itself. But clearly this is all nonsense, for these mistaken views invert the true relationship between these two things: architectural blueprints are only means to the ends which they help engineers and construction workers to realize, which is to construct, in accordance with the specifications contained in the blueprint, the edifice, bridge, road, or other human construction that it details artists’ paints are produced solely in order that painters can create paintings with them recipes are recorded, preserved, and propagated in order to help people prepare the dishes that they describe musical notes and scores are merely aids to musicians to help them play a composition accurately and letters exist in order that writers may convey their thoughts, beliefs, experiences, and theories – as well as their mistaken ideas and theories or invented fictions, in the case of writers like Dawkins – to others. [8]

Similarly, genes do not exist in, of, or for themselves: they exist so that they may give rise to the extraordinary living organisms that result from them, since genes are, in a very real sense, the blueprints of Life. In other words, Richard Dawkins has put the cart before the horse by calling all living organisms mere survival machines. Or, to use another analogy, Dawkins is like a man who takes a telescope and looks through the wrong end, all the while insisting that this is the right way to view the world. If anything, it is genes that serve the creatures to which they give rise, and not the reverse. Throughout his book, Dawkins has sought strenuously to objectify, belittle, and minimize all Life, which is alive. Concomitantly, he has sought just as strenuously to glorify a part of life that is not alive. However, I do not believe it is helpful to regard genes and the organisms they give rise to in this strict dichotomous manner, since this artificial division encourages people to regard organisms and their genes as separate entities, and therefore leads to the sorts of questions and distinctions that Dawkins is wont to pose or make in his book. Stated simply, to use Dawkins’ master/servant terminology in describing the relationship between genes and organisms is wrong and thus highly misleading.

There is another analogy that can help us to clarify the relationship between genes and the organisms they give rise to. Motors have many applications they power a variety of useful devices, including cars, fans, airplanes, lawn mowers, boats, electricity generators, and numerous other machines. However, even though motors can be separated physically from the devices which they power, and some of their parts can be replaced by other parts, no motor is built as an end in itself. Their function, their purpose, their raison-d’être can only be understood in relation to the devices that, in a very real sense, they animate. Similarly, although genes can be considered separately from the organisms they beget, and they can even be separated physically from these organisms, their function, purpose, and raison-d’être can only be understood in relation to these organisms. An important fact to remember about all genes is that, merely from examining a particular genetic sequence, it is impossible to know a priori what biological form, behaviour, or effect it will produce for this can only be discovered by examining the morphology or behaviour of the creature it gives rise to. To separate genes arbitrarily from the organisms they give rise to, as Dawkins adamantly insists that we must do, is completely to misunderstand their true nature and purpose. It is not the case, as Dawkins believes, that organisms arose solely for the purpose of enabling genes to reproduce and perpetuate themselves rather, the relationship between them is the reverse, for genes arose as a means of enabling organisms to reproduce and perpetuate themselves.

One way of sorting this whole matter out is to use the terms ‘replicator’ and ‘vehicle’. The fundamental units of natural selection, the basic things that survive or fail to survive, that form lineages of identical copies with occasional random mutations, are called replicators. DNA molecules are replicators. They generally, for reasons that we shall come to, gang together into large communal survival machines or ‘vehicles’. The vehicles that we know best are individual bodies like our own. A body, then, is not a replicator it is a vehicle. I must emphasize this, since the point has been misunderstood. Vehicles don’t replicate themselves they work to propagate their replicators. Replicators don’t behave, don’t perceive the world, don’t catch prey or run away from predators, they make vehicles that do all those things. For many purposes it is convenient for biologists to focus their attention at the level of the vehicle. For other purposes it is convenient for them to focus their attention at the level of the replicator. Gene and individual organism are not rivals for the same starring role in the Darwinian drama. They are cast in different, complementary and in many respects equally important roles, the role of replicator and the role of vehicle.

The replicator/vehicle terminology is helpful in various ways. For instance it clears up a tiresome controversy over the level at which natural selection acts. Superficially it might seem logical to place ‘individual selection’ on a sort of ladder of levels of selection, halfway between the ‘gene selection’ advocated in Chapter 3 and the ‘group selection’ criticized in Chapter 7. ‘Individual selection’ seems vaguely to be a middle way between two extremes, and many biologists and philosophers have been seduced into this facile path and treated it as such. But we can now see that it isn’t like that at all. We can now see that the organism and the group of organisms are true rivals for the vehicle role in the story, but neither of them is even a candidate for the replicator role. The controversy between ‘individual selection’ and ‘group selection’ is a real controversy between alternative vehicles. The controversy between individual selection and gene selection isn’t a controversy at all, for gene and organism are candidates for different, and complementary, roles in the story, the replicator and the vehicle. (254)

Dawkins declares, “Vehicles don’t replicate themselves they work to propagate their replicators.” But we do not need to swallow this morsel of sophistry, for we can declare instead, rejecting Dawkins’ arbitrary separation of organisms from their genes – a separation that is central to his selfish-gene theory, that the true function of genes is to enable organisms to replicate themselves. And this is, in fact, the commonsensical view that prevailed before the recent influence of Dawkins’ sophistical work, The Selfish Gene.

Dawkins goes on to declare, “The replicator/vehicle terminology is helpful in various ways.” However, he fails to recognize that it is also highly misleading, for it leads both him and his readers to think of genes and the organisms they give rise to as belonging to two fundamentally different categories of objects, which perform two fundamentally different functions, namely, replicating and preserving. In other words, the way that Dawkins defines these things paves the way for, and makes his readers more likely to accept, his pseudo-scientific theory.

Similarly, although it is possible to conceive of the heart, lungs, brain, or liver separately from the organisms in which they are found, and it is also possible to separate these organs physically from these organisms, no one has ever found a living heart, lung, brain, or liver existing in the natural world in complete separation from an organism. Hearts never developed to circulate, lungs to oxygenate, livers to filter, store, secrete, and eliminate, nor brains to command, control, perceive, and coordinate, outside or independently of the organisms of which they form an integral and essential part. For these vital organs exist only in relation to, within the bodies of, and for the sole purpose of helping to sustain large multicellular organisms. And, despite Richard Dawkins’ tiresomely insistent and incessant efforts to make us believe otherwise, neither did genes develop in order to exist independently of the organisms to which they give rise.

Since Dawkins has very clearly inverted the true relationship between organisms and their genes, by giving precedence to the second over the first, the primary criterion by which we should consider his theory is its ability to explain real-world phenomena, and whether it is able to explain these phenomena better than all rival theories. Judged by this criterion, it is not notably successful, for, as we shall see, although Dogmatic Dick Dawkins will never admit it, there exist many real-world behaviours that clearly contradict his theory, particularly in the realm of human behaviour.

The use of the word “machine” in Dawkins’ invented phrase “survival machine” reveals another aspect of Dawkins’ world-view, or what the Germans call a person’s Weltanschauung. To call all living organisms “machines” clearly indicates that Dawkins is fascinated by the development of man-made machines, robots, and computer technology. In other words, Richard Dawkins is, through and through, a technophile. Hence, it is not at all surprising that he has sought to formulate a wholly mechanistic explanation for the world and everything in it, including Life. Individuals like Dawkins believe that living organisms are merely the sum of their separate parts, and hence, there is nothing mysterious or miraculous about Life – meaning features that cannot be explained by science. But I completely disagree with this view, for although science, and biology in particular, has been highly successful in explaining Life’s numerous complex processes in physical and chemical terms, it still has not been able to explain why Life exists in the first place, and how a living organism can fashion itself out of inanimate elements such as carbon, oxygen, nitrogen, and hydrogen.

Most scientists and researchers, although they may be strongly attached to their particular beliefs, theories, or convictions, are ready to modify or abandon them if the evidence contradicts them. Moreover, they continually attempt to test their beliefs, theories, and hypotheses by comparing their predicted results with real-world data and observations. Let us consider the behaviour of Richard Dawkins whenever he is faced with real-world behaviours that contradict his selfish-gene theory.

Whales and dolphins drown if they are not allowed to breathe air. Baby whales, and injured individuals who cannot swim to the surface have been seen to be rescued and held up by companions in the school. It is not known whether whales have ways of knowing who their close relatives are, but it is possible that it does not matter. It may be that the overall probability that a random member of the school is a relation is so high that the altruism is worth the cost. Incidentally, there is at least one well-authenticated story of a drowning human swimmer being rescued by a wild dolphin. This could be regarded as a misfiring of the rule for saving drowning members of the school. The rule’s ‘definition’ of a member of the school who is drowning might be something like: ‘A long thing thrashing about and choking near the surface.’ (100)

Mistakes of this sort may, however, occasionally happen in nature. In species that live in herds or troops, an orphaned youngster may be adopted by a strange female, most probably one who has lost her own child. Monkey-watchers sometimes use the word ‘aunt’ for an adopting female. In most cases there is no evidence that she really is an aunt, or indeed any kind of relative: if monkey-watchers were as gene-conscious as they might be, they would not use an important word like ‘aunt’ so uncritically. In most cases we should probably regard adoption, however touching it may seem, as a misfiring of a built-in rule. This is because the generous female is doing her own genes no good by caring for the orphan. She is wasting time and energy which she could be investing in the lives of her own kin, particularly future children of her own. It is presumably a mistake that happens too seldom for natural selection to have ‘bothered’ to change the rule by making the maternal instinct more selective. In many cases, by the way, such adoptions do not occur, and an orphan is left to die. (101)

In the first of the two previous excerpts, Dawkins’ inordinately strong attachment to his theory leads him to make the ridiculous surmise – which, incidentally, as is true of many of his other declarations, he never bothered to verify – that this altruistic dolphin was not able to tell a human being apart from another dolphin! We could ask, How on earth was such a perceptually-challenged dolphin able to survive, since it would have had great difficulty distinguishing a fellow dolphin from a potential predator like a shark or killer whale? However, we need not venture into the marine world to find examples of inter-species altruistic behaviour, since there are many humans who go to great lengths to feed, save, raise, heal, nurture, or protect immature, sick, defenceless, or endangered members of other species. In most of these cases, it cannot be said that their altruistic behaviour is due to their desire selfishly to propagate their own genes, since these animals belong to a completely different species from them. For example, people who leave food for wild animals, such as in bird feeders, do so because they are fond of these animals and want to help them to survive, while having the opportunity to observe them when they come to feed. But although their actions may afford them pleasure or satisfaction, they do not in any way help their own genes’ survival, since it is not their intention to catch, eat, sell, or otherwise profit from the animal. In human communities where food is scarce, this sharing of food with non-human animals actually reduces the altruistic person’s chances of survival. In other words, these common human instances of altruism towards the members of other species very clearly contradict Dawkins’ selfish-gene theory. [9]

In both excerpts, Dawkins attempts to account for these altruistic behaviours that contradict his theory by cavalierly – and unscientifically – dismissing them as instances of the “misfiring of a build-in rule.” In the second excerpt, he declares, “In most cases we should probably regard adoption, however touching it may seem, as a misfiring of a built-in rule. […] It is presumably a mistake that happens too seldom for natural selection to have ‘bothered’ to change the rule by making the maternal instinct more selective.” First of all, Dawkins has simply invented this rule – that all biological mothers will only care for their own young – but without actually testing it, in order to dismiss, in this completely unscientific manner, this animal behaviour that poses a serious problem for his theory, since the raising of mammalian offspring often requires many years of devoted attention on the part of the parent. Secondly, in the human world, we know that there are many couples that have voluntarily adopted a baby from a different culture or ethnicity, and in all of these cases, it is clear that the baby does not share the parents’ genes, since it belongs to a different ethnicity from them. And even in cases where the baby is of the same ethnicity as the parents, there are again many instances where it is not related to them in the very narrow sense of Dawkins’ selfish-gene theory. Moreover, many of these parents must pay a large sum of money in order to adopt the baby, which represents a significant monetary sacrifice on their part, meaning that, far from gaining something from adopting the baby, they must forego the labour which they had to perform in order to earn that money, money which they could have used to raise – selfishly, according to Dawkins – their own biological children. I suspect that if these adoptive parents knew about Dawkins’ dismissive description of their altruistic behaviour – “as a misfiring of a built-in rule” – they would be highly incensed by it.

There is another serious problem with Dawkins’ theory: the genetic relationships that he works out in terms of the percentage of shared genes – 1 in the case of identical twins, .5 in the case of children and their parents, as well as between siblings of the same parents, .25 in the case of nieces, nephews, and grandchildren, and so forth – are invariant in both time and place. And yet, anyone who knows anything about different societies, as well as the significant changes that have occurred in many Western societies in recent centuries, knows that there exist significant differences between customary behaviours in different societies. Even the definition of “family” differs depending on the society one grows up in. In individualistic Western societies, this term is usually limited to what is called the nuclear family, which consists only of two parents and their offspring, and which is a relatively recent development that did not exist in the past. But in non-Western societies, the definition of family is more broad, and people are willing to help those, such as cousins, aunts, nephews, and so on, who would not be considered as “family” in the more individualistic and selfish West. In other words, except by resorting to ad hoc explanations – an activity at which Dawkins excels – meaning explanations that are invented expressly for the purpose of accounting for these significant variations and contradictory real-world behaviours, the selfish-gene theory simply does not allow for these common cultural variations in human behaviour, since, in genetic terms, a first cousin is a first cousin regardless of the society one grows up in, just as a sibling is a sibling, a biological parent is a biological parent, and so on in the case of aunts, uncles, and grandchildren.

In the case of humans, attachment results from spending a lot of time staring at, listening to, or otherwise perceiving something. In general, mothers spend much more time in close company with their children than fathers, which is one of the main reasons why mothers are more strongly attached to their children than fathers. Of course, the prevalent models of behaviour in a society are also important, for these will constrain people to do certain things, such as caring for one’s children or supporting them financially and materially, even in cases where one does not feel the urge to do so. These models also prevent people from caring for those who are not their own biological children. For example, there are many teachers, nurses, and others who are in regular contact with children and would like to do things for those children whom they become strongly attached to, such as buying clothes for them, which normally are performed only by parents. In these cases, the dominant models of behaviour will usually prevent them from acting on their generous impulses, lest other people misinterpret their behaviour or criticize it.

Humans do not only become attached in this way to their children, siblings, and other close relatives whom they see often. [10] They also become attached to other people, non-human animals, plants, articles of clothing, buildings, places, artistic creations, and other inanimate objects. In all cases where there is a strong attachment, there is a desire to preserve the person, place, building, animal, or object and protect it from harm. Such instances cannot be explained by Dawkins’ theory since, in the case of non-human animals, there exists no genetic relationship to account for the strong attachment. The same objection exists in the case of objects that do not in some way contribute to a person’s survival or likelihood of procreating and passing on one’s genes.

In addition, there are many cases of parents who have been separated at birth from their children, or siblings who have likewise been raised separately from birth or from an early age. According to Dawkins’ theory, these individuals should still exhibit the same concern for the welfare of the other person as exists in cases where there has been no separation, since the percentage of shared genes is the same in both cases, namely, fifty percent. But there are many instances where this is not true, including cases where the parties discover their genetic relatedness, and these examples also contradict Dawkins’ theory. In cases where a relationship develops between the two individuals, it is for the same reason that relationships develop in general: because the individuals spend time with each other and thus become fond of or attached to each other.

This belief was encapsulated in the statement, “Blood is thicker than water,” or “Blood will tell,” [11] that is, that the existence of a close genetic relationship, whether parent-child, brother-sister, or some other close familial relationship, would invariably determine the behaviour of both parties. But the many contradictory examples eventually discredited this belief. In a very real sense, what Dawkins has done is to revive this ancient, discredited belief by cloaking it in the modern language of genes.

In the excerpts about the dolphin and mammalian adoption, Dawkins is saying in effect, “The selfish-gene theory is correct it is the facts that are wrong. Any behaviours that seem to contradict my wondrous theory are either statistically insignificant or are merely examples of the misfiring of a rule that I will invent expressly for the purpose of defending my theory against its many detractors who cannot see that I am right.” In other words, Richard Dawkins will not admit that there exist many real-world behaviours that contradict his theory or cannot be explained by it. Despite his professions of scientific objectivity, Dawkins’ behaviour demonstrates that he is no different from the many dogmatic religious persons who refuse to admit that there exist many facts and much evidence that contradict their religious beliefs. For whenever he is confronted with a real-world behaviour that contradicts his theory, Dawkins dismisses it in this unscientific manner.

In the following excerpt, Dawkins appears to declare that facts cannot disprove or falsify a general theory, such as his selfish-gene theory:

The disagreement over lions [whether cooperative behaviour in lions is due to kin selection or reciprocal altruism] can be settled only by facts, and facts, as ever, tell us only about the particular case, not the general theoretical argument. (295)

And yet, in numerous other instances, Dawkins has no qualms in forming generalizations from specific facts, examples, or behaviours.

If an individual could be sure that a particular person was his identical twin, he should be exactly as concerned for his twin’s welfare as for his own. Any gene for twin altruism is bound to be carried by both twins, therefore if one dies heroically to save the other the gene lives on. Nine-banded armadillos are born in a litter of identical quadruplets. As far as I know, no feats of heroic self-sacrifice have been reported for young armadillos, but it has been pointed out that some strong altruism is definitely to be expected, and it would be well worth somebody’s while going out to South America to have a look. (93)

Had Dawkins behaved like a true scientist, he would have regarded the study of the nine-banded armadillo’s behaviour as an opportunity to test his selfish-gene theory. But after admitting that there is, as yet, absolutely no confirming evidence in this animal’s behaviour for his precious theory, Dawkins expects that its behaviour will conform to it. He again exhibits this unscientific attitude – that the facts will support his theory – in the following excerpt:

More generally, the Lack type of hypothesis is powerful enough to account, in selfish gene terms, for all evidence that might seem to support the group-selection theory, should any evidence turn up. (122)

In other words, Dawkins is saying that “any evidence that may be discovered in the future will support the Lack type of hypothesis, which I favour, since it is consistent with my selfish-gene theory, and will contradict the group-selection theory, which I despise.” This is truly an extraordinary claim on Dawkins’ part, for he is claiming that he has the ability to foresee the future!

As the last excerpt illustrates, Dawkins is extremely hostile towards any attempt to consider organisms in terms of groups, flocks, swarms, clusters, schools, herds, or human societies. This is because it is antithetical to the particular, Dawkinsian way of viewing the biological world, which is to focus one’s gaze unwaveringly on individual genes, only genes, and nothing but genes.

But we have once again slipped back into looking at life from the point of view of the individual organism rather than its genes. (250)

This packaging of living material into discrete vehicles became such a salient and dominant feature that, when biologists arrived on the scene and started asking questions about life, their questions were mostly about vehicles—individual organisms. The individual organism came first in the biologist’s consciousness, while the replicators—now known as genes—were seen as part of the machinery used by individual organisms. It requires a deliberate mental effort to turn biology the right way up again, and remind ourselves that the replicators come first, in importance as well as in history. (265)

The rivalry between individual organism and group of organisms for the vehicle role, being a real rivalry, can be settled. As it happens the outcome, in my view, is a decisive victory for the individual organism. The group is too wishy-washy an entity. A herd of deer, a pride of lions or a pack of wolves has a certain rudimentary coherence and unity of purpose. But this is paltry in comparison to the coherence and unity of purpose of the body of an individual lion, wolf, or deer. That this is true is now widely accepted, but why is it true? Extended phenotypes and parasites can again help us. (254-255)

Words like “wishy-washy” and “paltry” are expressions of Dawkins’ disdain towards this common view of, and approach to studying, animal behaviour. Depending on the context, Dawkins is dismissive of any attempt to consider organisms either as discrete wholes, as most of us consider them, or as members of a cooperating group. Repeatedly, as in the preceding excerpts, he harangues and exhorts his readers to regard organisms in exactly the same way that he regards them. In effect, Dawkins is saying that natural selection can only act either at the level of the individual gene or the individual or group, but not both. But this is a misleading either/or that seeks to reduce the complexity and heterogeneity of the real world to fit certain principles that may not accurately describe, or accord with, reality.

Like other dogmatic individuals, Richard Dawkins is wholly unaware of the distorting and deluding effects that one’s prejudices, whether they are enthusiastic or scornful, have on one’s perception of reality, and on one’s attitude towards a particular idea. In many cases, what he calls impeccable logic is nothing more than his inordinately strong desire to demonstrate to others that his views are correct. In considering the views of someone who has such strong convictions, it is important for us to consider impartially whether Dawkins’ strong enthusiasms, aversions, dislikes, and prejudices are in fact supported by the facts – and not merely the facts, as well as his personal interpretations of them, that he presents to his readers to buttress his views and theory.

As the next excerpt illustrates, Dawkins often mistakes the particular way that he views the world as revealing something important about the world.

I speculate that we shall come to accept the more radical idea that each one of our genes is a symbiotic unit. We are gigantic colonies of symbiotic genes. One cannot really speak of ‘evidence’ for this idea, but, as I tried to suggest in earlier chapters, it is really inherent in the very way we think about how genes work in sexual species. (182)

However, Dawkins fails to understand that this conjecture is nothing more than a particular way of perceiving, or categorizing, certain aspects of the world, a perspective that is not shared by others, except perhaps by those who admire him. When he writes, “it is really inherent in the very way we think about how genes work in sexual species”, what he really means is that this idea exists in the brain of Richard Dawkins, and therefore – even though, as he admits, there can never be any evidence for it – it must be true, since he is such a smart and clever person.

The next excerpt illustrates a curious recurring flaw in Dawkins’ understanding – or, more accurately, his misunderstanding – of reality:

To me, the most puzzling feature of naked mole rats is that, although they are like social insects in so many ways, they seem to have no equivalent caste to the young winged reproductives of ants and termites. They have reproductive individuals, of course, but these don’t start their careers by taking wing and dispersing their genes to new lands. As far as anyone knows, naked mole rat colonies just grow at the margins by expanding the subterranean burrow system. Apparently they don’t throw off long-distance dispersing individuals, the equivalent of winged reproductives. This is so surprising to my Darwinian intuition [12] that it is tempting to speculate. My hunch is that one day we shall discover a dispersal phase which has hitherto, for some reason, been overlooked. It is too much to hope that the dispersing individuals will literally sprout wings! But they might in various ways be equipped for life above ground rather than underground. (315)

Dawkins’ reasoning is as follows: because naked mole rats resemble social insects like bees and ants “in so many ways,” specifically the fact that, in naked mole rat colonies, there is only one reproducing female whose progeny the other mole rats help to raise, feed, and protect, it logically follows that they must also resemble social insects in other important respects, including the ants’ production of male and female reproductive individuals during a certain part of the year in order to found new colonies. In other words, according to Richard Dawkins, the Remarkable, Redoubtable Rationalist of Evolutionary Behaviour, things that resemble each other in some ways must also resemble each other in other important ways. But why must this be true? After all, despite their behavioural similarities, naked mole rats are also very different from bees and ants in many respects, such as the fact they have four legs instead of six, they are covered with skin instead of chitin, they have teeth instead of mandibles, they do not fly during any part of their life cycle, and they lack the antennae possessed by all social insects. [13] What this example shows is that Richard Dawkins expects reality to conform to his personal conceptions of it or to put it another way, Dawkins has mistaken his particular conceptions of reality for reality. Time and again throughout his book, we observe this Dawkinsian tendency manifested in one form or another, a tendency that, alas, often leads him into error.

Richard Dawkins is a scientific rationalist, since he worships at the Twin Temples of Reason and Science. However, among their many other shortcomings and proneness to error, the problem with dogmatic rationalists like Dawkins is that they assume that, in order to be true, a law, principle, or theory must be true without exception, in all times, places, and situations. But this naive and frequently erroneous philosophical belief overlooks the exceptions that exist to many generalizations, as well as to certain laws or theories such as Newton’s Law of Gravitation, or the Theory of Evolution by natural selection. I have considered these important exceptions to the latter theory in several other essays. [14] Like his avowed intellectual master, Charles Darwin, Dawkins has failed to see that the Theory of Evolution is incomplete because it examines life solely from the perspective of the individual organism, species, or gene – which latter is the unit that Dawkins dogmatically insists that we should narrowly view and reduce all Life. For when life is examined from the holistic perspective, a different pattern emerges, wherein cooperation rather than competition is most evident and important. But unlike Darwin, who wasn’t aware of the existence of genes, Dawkins believes that competition at the level of the gene is another invariable principle, which has led him to apply this principle to many situations where it is inappropriate or obviously wrong.

The following excerpt show that Dawkins believes that human behaviour is no different from animal behaviour, in the sense that most of our actions are determined directly by our genes, as is the case with the great majority of plant and animal behaviours.

Genetic relatives will tend to be alike not just in facial features but in all sorts of other respects as well. For instance, they will tend to resemble each other with respect to genetic tendencies to play—or not to play—Tit for Tat [a prisoners’ dilemma strategy, where each player does what the other player did in the last round]. So even if Tit for Tat is rare in the population as a whole, it may still be locally common. In a local area, Tit for Tat individuals may meet each other often enough to prosper from mutual cooperation, even though calculations that take into account only the global frequency in the total population might suggest that they are below the ‘knife-edge’ critical frequency. (219)

One might as well declare that, to paraphrase Dawkins, “In a local area, persons with a genetic predisposition to speak Spanish may meet each other often enough to prosper from speaking the same language, since the ability to speak a common language is a highly advantageous group trait.” But of course, no such genetic predisposition exists in the case of languages, and the fact that those who live in the same area very often do speak the same language is due to an entirely different process, namely, the tendency of human beings to imitate what they see and hear other humans doing, and their tendency to conform to the most common behaviours that are performed by those who are in their realm of influence. As this example shows, Dawkins has made the same mistake that Darwin made by assuming that most human behavioural characteristics are also genetically determined, just like their physical attributes.

In the following excerpt, Dawkins takes a moment from pontificating on his selfish-gene theory to give his readers a lesson in pronunciation:

Their [hydras’] tissues tend to be parasitized by algae. (The ‘g’ should be pronounced hard. For unknown reasons some biologists, not least in America, have recently taken to saying Algy as in Algernon, not only for the plural ‘algae’, which is—just—forgivable, but also for the singular ‘alga’, which is not.) (244)

As many foreign students of English have remarked, English pronunciation is frequently irregular, meaning it has many exceptions to its rules of pronunciation, unlike more regular languages like Spanish, Italian, and German, which are easier to pronounce. [15] For example, the English pronounce “lieutenant” as if it were spelled “leftenant.” And the middle letters in “Worcester” and “Gloucester” are not pronounced, so that they sound like “Wuster” and “Gloster,” as in the sentence, “My Lord, the noble Dukes of Wuster and Gloster have arrived.”

The rule that Dawkins is applying in this case is the rule that, when the consonants “c” and “g” are followed by the vowels “e” and “i,” they are pronounced softly, as in “gem,” “gin,” “celebration,” and “circle,” such as in the sentence “Although gems are reason for celebration, too much gin will make you think in circles” but when they are followed by the vowels “a,” “o,” and “u,” they are pronounced as hard constants, as in “cat,” “cold,” “cucumber,” “gave,” “gold,” and “guard.” An example is the nonsensical sentence, “The cat caught a cold from eating the cucumber, so it gave the gold to the guard to guard.” And yet, there are exceptions to this rule. For example, “sceptic” and “arcing” are both pronounced with a hard “c,” even though they are followed by one of the two softening vowels. To make this distinction clear, “sceptic” was formerly spelled “skeptic,” a spelling that has, however, largely become archaic. In the case of the letter “g,” “get,” “geyser,” “giddy,” “gimmick,” and “girl” are all pronounced with a hard “g,” even though the “g” is followed by an “e” or “i.” In the case of “get,” this is probably so that it remains recognizably consistent, in its spoken form, to the other conjugations of this irregular verb: besides “gets” and “getting,” these are “got” and “gotten,” all of which are pronounced with a hard “g.” If a person were to say, “Jet lost, you dick!” in accordance with the usual pronunciation of the combination “ge,” one would probably be able to understand the person’s meaning from his or her menacing expression and threatening tone of voice, but one might wonder if the person were drunk, had a speech impediment, or is a fervent disciple of Richard Dawkins.

More to the point, perhaps, the word “margarine” is pronounced with a soft “g” even though it is followed by an “a,” just like in “algae.” So instead of being pronounced like the “g” in “Margaret,” it is pronounced like the “j” in “marjoram,” which suggests the imperative, “Only marjoram-flavoured margarine for our dear Margaret!”

The plural “algae” is due to its Latin derivation, in which language the plural of a feminine noun is formed by adding an “e” to its singular nominative form. Hence, since most English speakers no longer speak Latin, there arose several different pronunciations of this irregular foreign word. The pronunciation that Herr Professor Dawkins criticizes – algy – in fact accords with the general rule of pronouncing a soft “g” when it is followed by the “e” sound – in this case, if not in its written form, then in its spoken form.

Besides showing that such irregularities in pronunciation do exist in the English language, there are several other reasons why I have spent some time considering what is admittedly a trivial aside. First, Dawkins uses the word “forgivable” – and implies its opposite, “unforgivable” – in describing the pronunciations of “algae” and “alga.” Clearly, he is highly irritated by those who, like me, pronounce “algae” as “alje.” Normally, one would not be motivated to use these words to describe what are, after all, minor variations or mistakes in pronunciation. This is not the only instance where Dawkins uses religious terms or metaphors in conjunction with his views, beliefs, and theory. In the following excerpts, he uses the word “orthodox” to describe his selfish-gene theory:

I have tried to do justice to Wynne-Edwards’s theory, even if rather briefly. If I have succeeded, you should now be feeling persuaded that it is, on the face of it, rather plausible. But the earlier chapters of this book should have prepared you to be sceptical to the point of saying that, plausible as it may sound, the evidence for Wynne-Edwards’s theory had better be good, or else. . . . And unfortunately the evidence is not good. It consists of a large number of examples which could be interpreted in his way, but which could equally well be interpreted on more orthodox ‘selfish gene’ lines. (115)

Frequently the evolutionary preconception in terms of which such theories are framed is implicitly group-selectionist, but it is possible to rephrase the theories in terms of orthodox gene selection. (191)

In other words, all those who dare to deviate from the “orthodox ‘selfish gene’ lines” are genetic heretics, at least in Dogmatic Dick’s view.

The following even more obviously Biblically-inspired sentence concludes a discussion of the fact that, in many bird species, the male is often colourful or possesses other decorative features, as is exemplified by the male peacock these features, although they may have evolved to attract the female, may also attract predators, or, by slowing them down, make it more difficult for the male to escape them: “What shall it profit a male if he shall gain the whole world, and lose his immortal genes?” (162) Dawkins’ meaning is that, although a drably-coloured and unremarkable male bird may be more likely to survive, it is also less likely to procreate and pass on its genes, since it will be overlooked or ignored by females. The original sentence occurs in a sermon preached by Jesus, as recorded in the Book of Mark, that exhorts his listeners to forsake worldly riches, pleasures, and even one’s life for the eternal spiritual rewards that await the faithful in the Kingdom of God.

For whoever would save his life will lose it, but whoever loses his life for my sake and the gospel’s will save it. For what does it profit a man to gain the whole world and forfeit his soul? (Mark 8:36)

Second, this example shows that Richard Dawkins is a dogmatic individual who wants to impose his views on everyone else, including the way they pronounce certain words like “algae.” [16] Throughout his book, Dawkins seeks to make people view the world in exactly the same way that he does. In addition, it shows that Dawkins does not like exceptions to general rules. In his ideal world, there would not be any exceptions to general rules, and certainly not to his precious selfish-gene theory, which he believes is true without exception. The problem with a theorist who is as dogmatic as Dawkins is that this tendency makes him blind to the many exceptions that exist to his theory or beliefs. In other words, Dawkins’ very strong preferences, beliefs, and prejudices make him an unreliable guide to finding out the truth about the world we live in – an important fact that a great many of his admiring readers have failed to recognize, since one effect of admiration is to make people uncritical of the admired person or object.

This dogmatic tendency is further illustrated by another book that Dawkins has written, called The God Delusion. Clearly, Richard Dawkins is deeply offended by the fact that so many people, in this scientific day and age, continue to believe in God. He is not content to live and let live, in the sense that, although he and many others do not believe in God, regarding it as a primitive, false, and possibly dangerous superstitious remnant of less enlightened times, he is intolerant of the many people who do believe in God, a belief that gives meaning, hope, discipline, and purpose to their lives. Although I believe in God, [17] and in fact I agree with him in believing that many of Christianity’s central beliefs, such as the belief in Mary’s immaculate conception, that Jesus was the son of God, and that devout believers are rewarded with eternal life in Heaven, while disbelievers are condemned to eternal punishment in Hell, are wrong, I am not bothered either by the fact that many people like Dawkins do not believe in God, or that many Christians and other religious people continue to believe things that I believe are wrong.

Dawkins exhibits considerable hostility to all religious beliefs, in no small part because they are rivals to the orthodox scientific account of the history of the world, which he obviously favours. It is, therefore, more than a little ironic that what Dawkins has done is to displace God the Creator in the typical creation myth with the Gene: whereas Christians believe that, In the beginning was the Word, as spoken by God, which Word created the World, Dawkins the scientific rationalist believes that, In the beginning was the Gene, which created and gave rise to all Life. Furthermore, just as religious people believe that everything in God’s Creation is subservient to God, he claims that all life forms are in reality subservient to their Genes, which are the true Masters of Life. And just as religious people attribute immortality to God, Dawkins attributes immortality to the Gene. As he explains in his book, an alternative title for it was The Immortal Gene. I suspect this is another reason why his book has aroused such hostility in many readers: Dawkins behaves as a sort of autocratic scientific inquisitor who seeks to eradicate all non-scientific – meaning primarily religious – heresies from the face of the Earth, in order to propagate the Cult of the Immortal Gene. This is illustrated by the venomous intolerance with which he condemns all those who dare to criticize his purportedly scientific beliefs.

There are other reasons why many people find Dawkins’ thesis – that all living organisms are merely the individual genes’ way of perpetuating themselves – objectionable. First of all, it attributes to a molecule, or a sequence of molecules, characteristics such as the desire to survive and perpetuate itself, which most people would attribute only to living organisms. This brings us to another very curious feature of Dawkins’ purportedly scientific theory: whereas science has sought assiduously to remove teleology, or the study of primal causes and ultimate ends, from its various disciplines, Dawkins has reintroduced it in the realm of biology. In other words, Richard Dawkins has made the mistake of attributing purpose to inanimate processes: because genes replicate, and some of them are able to perpetuate themselves for long periods of time, this means that this is what they intend or strive to do.

In the past, and even today, many people attributed intentions, desires, and feelings to inanimate phenomena such as lightening, thunder, rain, earthquakes, tides, volcanic eruptions, the changing of the seasons, the rising and setting of the Sun, the moon’s waxing, waning, and occasional eclipsing, and so forth. Even the great Aristotle attributed the desire to do certain things to inanimate objects, such as that moving objects come to rest because they seek to do so. These were common beliefs prior to the development of the modern scientific outlook. Similarly, by claiming that genes are selfish, Dawkins has attributed a teleological goal to their behaviour, which most people would regard as ordered but nevertheless inanimate processes. What Dawkins has done is to shift all the wonder and mystery that most people feel about Life and redirected it to our genes. According to Richard the Genetic Animist, since genes give rise to all life forms on the Earth, it follows that they themselves must be alive.

Dawkins’ behaviour is extremely curious, for, like a scientific rationalist vampire, he has drained all the life out of Life, by declaring that living organisms are merely empty survival vehicles for their genes, and then transferred this animating force and purposive volition to genes, declaring that they are in fact immortal, even though genes are not alive. Earlier, I observed that Dawkins has probably read a considerable amount of science fiction. A common theme in this genre is an alien life-form that deviously and ruthlessly enslaves the human race for its own purposes, such as to ensure its own survival. Without pursuing it further, I will make the assertion that Dawkins’ selfish-gene theory shares many similarities with this kind of writing, which has as its central theme the possession, control, conquest, invasion, infiltration, or contamination of terrestrial organisms by an alien life-form solely for its own selfish and nefarious purposes. [18]

The majority of people in the world have not read Dawkins’ book. But there are many people who have heard the phrase, “the selfish gene.” What meaning does this phrase suggest to them? In my opinion, it leads them to believe that our genes make us behave selfishly and since a person’s genes cannot be changed, this means that we cannot help behaving selfishly – in other words, that selfishness is an innate human behavioural trait. However, unlike Dawkins’ narrow technical definition of selfishness, which is “whatever is for the good, survival, and multiplication of our genes,” most of those who have not read but have heard of The Selfish Gene probably take it to mean “whatever is good for me, and perhaps those few people I care about.” There are other sources of this mistaken belief, notably economics, which assumes that human beings are rational and selfish maximizers of their personal utility but Dawkins’ book is another source, and one whose influence, unintended though it may be, has not previously been recognized in areas outside of its specific area of intended applicability. [19] In other words, whereas the free-market economists have declared that all people should behave selfishly, for this course will, at the same time, also help to promote other people’s welfare and well-being, Dawkins book, or at least the popular impression of it, declares that we cannot help behaving selfishly.

The highly confrontational consequences of Dawkins’ mistaken theory are shown by the following excerpt: “The selfish-herd model in itself has no place for cooperative interactions. There is no altruism here, only selfish exploitation by each individual of every other individual.” (168) Dawkins even extends this selfishness to relations between parents and their offspring, since, in the case of organisms that reproduce sexually, a parent and child share, on average, only 50% of their genes, since the other 50% of the child’s genes were contributed by the other parent, and hence, these genetic differences will cause them to compete selfishly for the survival of their particular genes:

There is really only one entity whose point of view matters in evolution, and that entity is the selfish gene. Genes in juvenile bodies will be selected for their ability to outsmart parental bodies genes in parental bodies will be selected for their ability to outsmart the young. (137)

Although Dawkins does not explicitly say so, it appears that he is attempting to account for the frequency of parent-child conflicts and disagreements in modern societies. But the true cause of these conflicts is the fact that, today, many children observe different models of behaviour than the models their parents grew up observing, a development that, in historical terms, is very recent. In the past, before modern methods of communication and the dominant Platonic method of education, which rigidly herds children into age-cohorts, the models observed by parents and their children were more or less identical, and so parent-child conflict did not exist back then, and even if it did, then it did not exist to the extent that it does today. Even today, in societies where people of different generations observe the same models of behaviour, there is much less intergenerational conflict than in societies where this is not the case.

In practical terms, the effect of Dawkins’ argument has been to promote and encourage people to behave selfishly by making them mistakenly believe that selfishness is an innate, meaning genetically determined, human trait, and therefore they can do little or nothing to change it. But besides the other examples we have considered of fairly common altruistic behaviours, if Dawkins’ theory is true, how, then, can we account for the behaviour of someone like Mother Teresa, who helped a great many people to whom she was not genetically related, except in the sense that they belonged to the same species as her? In her case, it was obvious that she was not helping those who were related to her, since she, a white Albanian woman, spent most of her adult life in India, where she helped numerous people who had a different skin colour and other differentiating physical traits from her. Many of these people, such as abandoned babies, would not have survived without her help and care. According to Dawkins’ theory, such exemplary examples of disinterested generosity should never occur, since they involve the total abnegation of the survival of one’s own genes, while one devotes oneself to helping other people’s unrelated genes to survive. And yet, Mother Teresa is by no means the only person who has devoted her life to helping people who were not genetically related to her, and without receiving any financial remuneration for her work.

None of the important facts in the life of Mother Teresa can be explained by Dawkins’ selfish-gene theory, for, as we shall see, these facts directly contradict it. Mother Teresa had no issue, and this was not because she was infertile, but because she took a vow of chastity, in imitation of Jesus and the many others, both nuns and monks, who likewise had no issue because of their desire to live as Jesus and their fellow nuns or monks did. The many millions of people who, throughout history, have taken a lifelong vow of chastity provide a real-world refutation of the selfish-gene theory, since the first necessary condition for propagating one’s genes is to have children. Hence, this behaviour cannot be dismissed pseudo-scientifically as the “misfiring of a built-in rule,” as Dawkins is wont to declare whenever he is faced with a behaviour that contradicts his theory, since these people all decided deliberately not to have any children, to the grave detriment of their genes’ survival. If Dawkins’ theory is correct, the question we need to ask is, “Why didn’t Mother Teresa’s selfish genes make her break her vow of chastity and beget children?”

Mother Teresa had an older brother and sister. Although her sister did not have any children, her brother had one son. In addition, she probably had cousins and other extended family members who had children. Again, based on Dawkins’ selfish-gene theory, we must ask ourselves, “Why, after she decided not to have any children of her own, didn’t Mother Teresa’s selfish genes make her remain in, or return to, her native Macedonia [20] and care for those who were related to her, thus ensuring that at least some of her genes would survive, rather than going to a foreign country and helping the genes of a great many foreigners, none of whom were related to her in the narrow biological sense, to survive?”

Clearly Mother Teresa was inspired by Biblical stories like the one about the Good Samaritan who helped a stranger who was robbed and assaulted by thieves, as well as by Jesus’s teachings and exhortations to love all human beings (but not all living creatures, which clearly shows the limits of Jesus’s sympathy and understanding). Moreover, she was also inspired by the example of missionaries who left their native country and went to foreign places in order to help those who lived there, where it was much less likely that the inhabitants would share their genes. These and many other examples of generosity towards those who are not genetically related to us very clearly contradict Dawkins’ dogmatic selfish-gene theory.

Albert Schweitzer is another prominent person who was somehow able to resist the irresistible urge of his selfish genes to care only for his own children, or those of his close relatives. Although he, unlike Mother Teresa, married and had one child, he and his wife decided that they would devote themselves to helping people in Africa, in part to atone for the ravages and horrors caused by European colonialism. To achieve this end, Schweitzer trained to become a doctor, in which role he saved or prolonged the lives of many Africans, who were thereby enabled to pass on their selfish genes, at the partial cost of Schweitzer’s and his wife’s own selfish genes, since they were not able to beget and care for more of their own children. Today, there exist numerous organisations, such as Médecins sans frontières, whose primary goal is to save the lives of as many people as possible in poorer countries in the world, or those who have been injured as a result of violence. [21] And the people who work for these organisations do not do so because of reciprocity, for in most cases they receive nothing from the people they help except perhaps their gratitude.

Another important point shown by these examples is that Richard Dawkins has fundamentally misunderstood the way in which genes operate. By attributing to them conscious or deliberate motives, specifically to ensure their own survival and increase their number, he has made the mistake of attributing purposive goals to non-living elements. Genes determine both the morphology and behaviour of many organisms [22] however, this does not mean that they also seek to ensure their own survival. A storm may cause considerable damage to the environment, both living and non-living, but no one would therefore conclude that the storm intended to cause all this harm, since we know that storms are not living things. The Sun shines on the Earth and, with a minute fraction of the total radiation it emits, thereby enables Life on Earth to continue to exist but no one, apart from a Sun-worshipper, would say that it intends to do this, since, first of all, the Sun is not a living entity, and second, there are many stars in the Universe, probably the great majority of them, whose radiation does not support any life.

The fight or flight response possessed by many animals helps them to survive by making them either aggressively defend themselves against predators and adversaries, or frantically try to escape from them. However, as the domestication of numerous animals species has shown, in which these natural responses have been either muted or eliminated, [23] these genetic behavioural responses are separate from the rest of the animal’s genes – those genes that determine the animal’s physiognomy, organs, and other physical characteristics. Similarly, it is also possible to imagine an animal that either lacks reproductive organs or, possessing these organs, lacks the mating behaviour typical of that animal. [24] Again, these organs and mating behaviour are completely separate from the rest of the animal’s genes but it is these genes that enable the animal to pass on all of its genes to future generations, or at least half of them to any given offspring. Hence, these highly useful evolutionary behaviours benefit not only these genes but also the rest of the animal’s genes. This is a clear example of how Dawkins’ monomaniacal insistence that we should only consider all behaviours from the very narrow perspective of the individual gene leads him and his many admirers into error and makes them overlook some pretty obvious truths about the world.

These examples show that, contrary to Dawkins’ belief that all genes selfishly compete with each other for their individual survival, genes do in fact cooperate and coordinate with each other. The genes that control for the fight or flight response, as well as the genes that control for the develop of the reproductive organs and mating behaviour, presuppose the existence of other genes that control for muscular, skeletal, visceral, neuronal, and other of the animal’s physical traits and abilities. Without the former genes, the latter genes either would not survive or would not be able to reproduce themselves but without the latter, the former would have no animal structure on which to act. Hence, contrary to Dawkins’ reductionist thesis, these genes act harmoniously together to ensure the survival and reproduction of the entire organism. [25]

Before Darwin presented the Theory of Evolution to the world, most people believed that God was responsible for directly creating and animating all living things, in the guise of a master puppeteer or divine magician: Life exists because God wills it to exist, and all living creatures are thus manifestations of God’s volition. But the Theory of Evolution showed that this common belief is not true. It is a quirk of Dawkins’ personality and upbringing that, instead of simply accepting that, as the Theory of Evolution declares, all Life is the result of accidental circumstances, specifically of the non-volitional, or unintended, results of the Law of Natural Selection, he must needs reintroduce purpose in the biological realm by transferring it from God to organisms’ genes. And this is due, I suspect, to the fact that, until he was a teenager, Dawkins believed in God and the Christian story of how the world was created. [26] For a person who had been raised as an atheist, receiving no instruction in any particular religion, would, unlike Dawkins, have felt no inclination to attribute purpose anywhere in the operations of the Universe, including the operations of Life on Earth, since one would conclude that these results are all due to the impersonal laws that have been discovered by science. But because Dawkins was accustomed to worshipping God, when he ceased believing in God, whether he was aware of it or not, he sought to replace the God-figure with a new idol that he could worship, which in his case was the Almighty Gene.

As we saw previously with the naked mole rat, where he mistakenly sought to impose another species’ behavioural repertoire on it, although, as a teenager, Dawkins rejected the religious, and specifically the Christian, explanation for the Universe and everything in it, he retained the structure of this religious model and unwittingly imposed it on his later study of genes and their relations to organisms. In the case of the naked mole rat, Dawkins’ prior knowledge of social insect behaviour interfered with his ability to understand the similar, but also dissimilar, behaviour of this animal. In the case of genes and organisms, however, the effect has been disastrous, for the religious model that he imbibed in his childhood has greatly distorted his understanding of the true relationship between genes and organisms. Moreover, the dogmatic tendency of many religious believers has led Dawkins, first, to impose a religious-style, pseudo-scientific uniformity on the world second, to a strong desire to make others conform to his views and third, to an egregious blindness to facts that contradict his precious theory.

If we examine the wondrous traits that Dawkins attributes to genes, as well as the reverential and deferential way that he speaks of them, we will see that his attitude towards genes shares many similarities with the way that religious people speak of God. Just as religious people believe that God is immortal, Dawkins believes that our genes are likewise immortal just as religious people abase themselves in relation to God, declaring that God is everything and we, Its humble creatures, are nothing, Dawkins abases all living organisms in relation to their genes, declaring that genes are everything and that we organisms are merely their temporary “survival vehicles” just as religious people believe that God uses – and some of them fervently desire God to use – them to achieve Its divine ends, Dawkins believes that our genes use us and all other organisms to achieve their selfish ends, which are to ensure their survival while multiplying themselves to the greatest possible extent just as religious people believe that God is timeless and exists independently of all Its living creations, Dawkins believes that the immortal replicators also exist independently of Life just as religious people believe that God’s presence is visible everywhere, Dawkins believes that the effects of the selfish gene are also visible everywhere there is life in the Universe and just as religious people believe that God existed prior to the creation of life, Dawkins believes that the immortal replicators also existed prior to the appearance of life and its subsequent evolution. What this shows is that Richard Dawkins is a scientific worshipper who has revived a form of animism – genetic animism – by prostrating himself, and, moreover, by wanting the rest of us to prostrate ourselves, before an inanimate entity, namely, our genes.

It should now be evident what Richard Dawkins’ ulterior motive is in presenting his selfish-gene theory: whether he realizes it or not, his aim is not primarily to discover the truth, but to promulgate a – in his view – logically consistent and comprehensive scientific theory that will displace the many religious, and specifically Christian, accounts, which have been believed and continue to be believed by many people, of the origin of life on Earth. In other words, his aim is no less religious than it is scientific, for in his writings, Dawkins exhibits the proselytizing zeal of the evangelist, albeit one clothed in scientific garb. This explains why he is so adamant in defending his precious theory, and why he will not admit the obvious fact that there exist many exceptions to it, for he is motivated by the same religious fervour that animates those who preach religious doctrines. If truth were indeed Dawkins’ primary concern, as it is for any objective scientist, then he would not behave in the prevaricating, pseudo-scientific way that he does whenever he considers examples that contradict his precious theory, by simply dismissing them as unimportant instances of the “misfiring of a built-in rule.” To Dawkins, truth is a secondary concern to defending and propagating evolutionary doctrine against religious people in general, and in particular against religious fundamentalists who believe in the literal truth of the Bible. He is as determined as they are to prove that his views, and not theirs, are correct. In other words, Richard Dawkins is a true scientific believer, and his book, The Selfish Gene, is a work of scientific theology.

The phenomenon of cults and religions shows that charismatic, egocentric, or domineering individuals can persuade others of things and inculcate practices that are wrong, bad, or harmful. For the truth is that most people are meek followers rather than strong-willed leaders, and therefore they are prone to following an individual who is able to persuade them by one’s charm or charisma, the conviction of one’s rhetoric, the force of one’s personality, or by some other, non-rational trait. And in spite of its usual veneer of impartiality, science is not immune to the influence of such individuals. Generally speaking, those who succeed are often individuals who are assertive, supremely confident in their abilities or that they are right, and so on.

I wish to introduce a concept that I will call the “Napoleon Syndrome.” [27] Napoleon Bonaparte was a person who was strong-willed, possessed extraordinary abilities and energy, and believed unquestioningly in his destiny to accomplish great things during his lifetime. Such individuals can be found in all fields of human endeavour, whether in politics, war, business, industry, science, technology, the arts, sports, and even romance. Many of humanity’s accomplishments are due to such individuals and their unshakable faith in themselves, as well as their indomitable desire to succeed. But these sorts of strong-willed individuals do not accomplish only good, for there are also individuals, such as Hitler, Stalin, Lenin, and Mao, who cause considerable harm by their forceful character and their unwavering belief that they are destined to accomplish great things. Because they are followed blindly and devotedly by their many admirers, they often lead their admirers and others down the path to ruin and disaster. These individuals have been described in various ways, such as “a force of nature,” or “those who carrying all before them like a flood” and the adulation with which they are regarded by others has been described as “hero-worship” or a “personality cult.” Some other common features of the Napoleon Syndrome are a tendency to be dogmatic, to expect unwavering loyalty and obedience from others, whom one generally regards as being inferior and subservient to oneself, to criticize severely, argue vehemently with, and condemn those who do not agree with one’s views, and to regard oneself as being above, or not constrained by, ordinary rules, laws, and social conventions – in other words, that they are a law unto themselves.

An important reason why these individuals are able to accomplish so much more than ordinary people is because they are able to gain other people’s steadfast admiration for themselves. Hence, when admiration is transmitted through a group of people by imitation, as it was among the French people for Napoleon in the last decade of the eighteenth and the early years of the nineteenth century, it enables these individuals to accomplish the extraordinary feats, whether good or bad, for which they are remembered, and for which their names are inscribed in the pages of history.

Following the deposition and later execution of the French King Louis XVI, which acts were in no way Napoleon’s doing, a war of some sort was likely between the newly christened French Republic and the other countries and regions of Europe that were still governed by hereditary monarchs. However, it is hard to imagine that this war would have lasted as long as it did, and caused the deaths of so many individuals, if someone other than Napoleon had gained control of, first, the French army, and later the French Republic, which he restored to its former monarchical form with himself as its head. [28] For the same military abilities that enabled him to gain the respect and reverence of both the French people and his soldiers also enabled him to remain in power much longer than a less able individual. Hence, without Bonaparte at its head, it is likely that France would not have been able to continue fighting against the countries that were allied against it for as long as it did.

In the less tumultuous and bloody realm of science, a recent example of the Napoleon Syndrome is Sigmund Freud. As more and more people have realized, Freud’s theory of human behaviour is complete nonsense. [29] And yet, for close to a century, there were many millions of people around the world who were convinced of its truth, and many patients who were willing to pay large amounts of money to be psychoanalysed, in the expectation of being cured of their neuroses, repressions, and other emotional problems and discontents. Since Freud’s theory of human behaviour is wrong, we can only explain his success in convincing others of its truth by the fact that he possessed many of the traits of the Napoleon Syndrome: a strong-willed character, the unwavering conviction that his theory was correct and that he was destined to accomplish great things in life, and intellectual abilities that greatly impressed those who either came in contact with him or read one or more of his many works.

A more distant historical example of the Napoleon Syndrome is Socrates, who, although he failed to compel his fellow Athenians to do his bidding, was nevertheless able, through the writings of his adoring pupil Plato, to influence many millions of people long after his death. Despite his professed modesty, Socrates was in reality an arrogant and conceited individual who had an unshakeable belief in his abilities and beliefs, while he incessantly denigrated the abilities and beliefs of others. Only someone who was as strong-willed and as sure of himself as Socrates was could have continued to question and publicly humiliate the prominent inhabitants of the city in which he lived, [30] when he surely must have been aware of the harmful effects of his incessant efforts to show that they knew nothing, namely, that he was gaining many prominent enemies and detractors by his unwise actions.

I have presented this digression because, in my opinion, Richard Dawkins belongs in the category of individuals who exemplify what I have called the Napoleon Syndrome, since he too is a domineering individual who is extremely sure of himself. Like Freud’s theory of the unconscious, Dawkins’ selfish-gene theory is wrong, or it is not as globally valid as he and his many admirers believe it is. [31] Hence, the only way to account for the fact that, in spite of its numerous flaws and inversions of the truth, its inability to account for many real-world behaviours, and the many behaviours that contradict it, his theory has been so widely accepted is by considering the character traits of its progenitor. It is by no means an exaggeration to call Dawkins the self-appointed High Priest of Natural Selection, to whom all others must defer. This would explain why Dawkins has been so dismissively critical of Stephen Jay Gould, since Dawkins has sought to supplant Gould, whom he regarded as a powerful and highly regarded rival, as the undisputed authority on, and champion and defender of, the Theory of Evolution.

There is another important similarity between Dawkins’ selfish-gene theory and Freud’s theory of the unconscious mind: in the way that they have been used by their originators, both these theories enabled individuals to attribute all manner of causes and explanations for human behaviours, in the case of Freud, and human and non-human behaviours, in Dawkins’ case. In Freud’s case, if a person objected that one was not aware of being motivated by, for example, the Oedipus or Electra complex, or the even more ridiculous concepts of penis envy and castration fear, Freud would have replied, “Aha, that is because, being unconscious motives, of course you are not aware of them. These primal motives can only be known by the effects they produce, which effects are uniformly bad when they are repressed. And that is why you must be psychoanalysed by me, my good man (or woman).” Similarly, throughout his book, Dawkins simply assumes that the genes for this or that behaviour must exist because, according to him, all behaviours are the direct result of our genes, which generally make organisms behave selfishly, except when it is in their genes’ selfish interest to make them behave altruistically.

Just as, in the case of human behaviour, to think of all people being motivated either by the Oedipus or Elektra Complex is the wrong way to think about human behaviour, in the case of biology, to think of genes selfishly trying to replicate and multiply themselves in successive generations of “survival vehicles” is the wrong way to think about genes.

The possessor of an altruistic gene might be recognized simply by the fact that he does altruistic acts. A gene could prosper in the gene pool if it ‘said’ the equivalent of: ‘Body, if A is drowning as a result of trying to save someone else from drowning, jump in and rescue A.’ The reason such a gene could do well is that there is a greater than average chance that A contains the same life-saving altruistic gene. (89)

But since Dawkins has not bothered to look for it, how can he be so sure that such a gene exists? There are many people who speak a specific language, such as Hindi, Swahili, or Portuguese, play a musical instrument, write novels, cook fancy dinners, sew clothes, make automobiles, ride bicycles, program computers, wear designer clothes, take photographs, hoard money, drink alcohol, play games, and smoke cigarettes but there are no specific genes that make people do any of these things, since these are all learned behaviours. Hence, since Dawkins has not actually taken the time and done the tedious work to find these altruistic genes, his belief that both altruistic and all other kinds of behaviour must be genetically determined is completely unfounded, since most human behaviours are not determined in this manner. This Freudian-like attribution of unconscious motives to people’s behaviour is even more evident in the next excerpt:

This chapter, and the next in which we discuss conflict between mates, could seem horribly cynical, and might even be distressing to human parents, devoted as they are to their children, and to each other. Once again I must emphasize that I am not talking about conscious motives. Nobody is suggesting that children deliberately and consciously deceive their parents because of the selfish genes within them. And I must repeat that when I say something like ‘A child should lose no opportunity of cheating . . . lying, deceiving, exploiting . . .’, I am using the word ‘should’ in a special way. I am not advocating this kind of behaviour as moral or desirable. I am simply saying that natural selection will tend to favour children who do act in this way, and that therefore when we look at wild populations we may expect to see cheating and selfishness within families. The phrase ‘the child should cheat’ means that genes that tend to make children cheat have an advantage in the gene pool. If there is a human moral to be drawn, it is that we must teach our children altruism, for we cannot expect it to be part of their biological nature. (139)

Just as Sigmund Fraud declared that many of our actions result from unconscious desires, Dawkins is saying that much of human behaviour is also due to unconscious motives, which are all due to our immortal genes’ selfish desire to propagate and multiply themselves in the world, regardless of what we, their humble and subservient “survival vehicles,” may want or intend to do.

At the end of his book, Dawkins summarizes his highly personal reinterpretation of Darwin’s Theory of Evolution in typically bombastic Dawkinsian style:

Let me end with a brief manifesto, a summary of the entire selfish gene/extended phenotype view of life. It is a view, I maintain, that applies to living things everywhere in the universe. The fundamental unit, the prime mover of all life, is the replicator. A replicator is anything in the universe of which copies are made. Replicators come into existence, in the first place, by chance, by the random jostling of smaller particles. Once a replicator has come into existence it is capable of generating an indefinitely large set of copies of itself. No copying process is perfect, however, and the population of replicators comes to include varieties that differ from one another. Some of these varieties turn out to have lost the power of self-replication, and their kind ceases to exist when they themselves cease to exist. Others can still replicate, but less effectively. Yet other varieties happen to find themselves in possession of new tricks: they turn out to be even better self-replicators than their predecessors and contemporaries. It is their descendants that come to dominate the population. As time goes by, the world becomes filled with the most powerful and ingenious replicators. (264-265)

But the individual body, so familiar to us on our planet, did not have to exist. The only kind of entity that has to exist in order for life to arise, anywhere in the universe, is the immortal replicator. (266)

In other words, according to Dogmatic Dick Dawkins, all that matters in the case of Life are DNA and the particular creatures which they give rise to are completely immaterial, insignificant, and unimportant, except as vehicles for perpetuating their Immortal Genes.

Incredibly, Dawkins declares that his selfish-gene theory is true of all life in the Universe, even though we have, until now, not discovered any life outside of our planet, which means that no one has yet been able to observe or study the different forms of life that may exist elsewhere in the Universe. To make such an outlandish, unwarranted, and unscientific generalization of his theory, which, as we have seen, is not even globally valid for all the living creatures that exist on the Earth, is incontrovertible evidence of Dawkins’ truly astonishing arrogance. As this excerpt clearly demonstrates, Richard Dawkins is an arrogant, deluded fool who has mistaken the contents of his brain for reality. But there is good reason for his many critics, detractors, and the subjects of his bullying to rejoice, for the intellectual reign of Richard the Rational, also known as Dogmatic Dick Dawkins, the animistic worshipper of the Immortal Gene, is well and truly over.

[1] The Selfish Gene by Richard Dawkins, p. 18. Oxford University Press, Oxford, 2006. Henceforth, as all the excerpts in this essay are taken from this book, only the page number will be cited immediately following the excerpt.

[2] In making this claim, I am using Dickens to represent fictional writers in general, and not to argue that Dawkins was influenced specifically by the works of Dickens. From a number of allusions and speculations, it is evident that Dawkins enjoys reading science fiction. For example, in The Selfish Gene, he spends several pages discussing an analogy taken from a work of science fiction that has something to do with the Andromeda Galaxy.

[3] I have considered the fallacious question “Which came first, the chicken or the egg?” in the essay “The Chicken and the Egg.”

[4] In the case of criminal investigations, investigators seek to find out the perpetrators of crimes. In attempting to reconstruct the actual course of events, they often must have recourse to indirect evidence and the testimony of witnesses, since they themselves were not there to observe the events firsthand. But it often happens that many a plausible case of how and why a certain accused individual committed this or that crime turns out to be wrong, thus leading to wrongful convictions and, on occasion, the execution of innocent persons, based merely on a probable or plausible account of how the crime was committed. Similarly, in science, there are many plausible theories that have been believed to be true for varying periods of time, until either they were shown to be wrong or they were displaced by another theory.

[5] This is no different from a person who, while constantly declaring that one has no intention of trying to get you to smoke, repeatedly offers you a cigarette and a light, while one smokes incessantly or a person who, while assuring you that one has no intention of trying to get you to drink, repeatedly places a glass of alcohol in front of you while consuming glass after glass of the stuff or a lusty man who, while declaring that one’s intentions are entirely honourable, repeatedly tries to seduce the woman of his desires.

[6] This is the famous play within the play “wherein I’ll catch the conscience of the King”, meaning Hamlet’s usurping Uncle Claudius, who killed Hamlet’s father and then wedded, and bedded, his mother.

[7] Despite Dawkins’ declaration that the title of his book is merely a useful analogy that is not to be taken literally, the fact is that, repeatedly throughout his book, Dawkins himself does take it literally – which literal interpretation leads him frequently into error – in the sense that each and every individual gene strives selfishly to protect and propagate itself as widely as possible. Similarly, prior to his book, it was commonly believed that individual organisms behave in this manner, by, for instance, defending themselves from predators, seeking selfishly to gain as much nourishment as possible for themselves and their offspring, and seeking to mate with as many members of the opposite sex as possible.

[8] Lest the assiduously argumentative and doggedly disputatious Dick Dawkins object that these are all examples of deliberate, that is, purposive human activities, and natural selection, in contrast, is an inanimate process, I declare that this is the very crux of our disagreement. For, like many another scientific rationalist or atheist, whereas Dawkins claims – at least when he is not attributing purposive characteristics like selfishness to genes – that all Life arose entirely by the concatenation of fortuitous accidental circumstances, in accordance with natural laws like the Law of Natural Selection, I categorically deny this belief.

[9] Of course, there are many instances where humans take care of animals primarily because they are useful to them in some way, whether by providing them with food, milk, protection, clothing, labour, transportation, or money. In these cases, their behaviour is clearly not motivated by altruistic motives.

[10] There are also many close blood relations, including parents and children, or brothers and sisters, who dislike or hate each other, and thus are unwilling to do anything to help their near relative. All of these fairly common instances, which again clearly contradict Dawkins’ selfish-gene theory, are due to contempt, an important determinant of human behaviour that Dawkins never considers.

[11] Another sense in which this phrase was meant was the presumed inheritance of certain characteristics, personality traits, or behavioural tendencies.

[12] What Dawkins mistakenly calls his “Darwinian intuition” is in reality his all-too-frequent proneness to Dawkinsian delusion, a tendency that Darwin himself did not suffer from, since he did not often make the mistake of trying to impose on facts an incorrect interpretation of them or an alien structure, as Dawkins does in this instance.

[13] Although all ant species live in colonies, there are many bee species that are individualistic, reproducing like most other sexual organisms, in pairs rather than in colonies or hives with only one egg-laying individual. Hence, even in the realm of social insects, there are many species that do not conform to the behavioural model that Dawkins attempts to impose on the naked mole rat’s behaviour.

[14] See “The Lion and the Lamb,” “Darwin’s Blunder,” “The Evolutionary Conundrum of Human Behaviour,” “Natural Selection Does Not Tell the Whole Story,” and “The Randomness Principle.”

[15] Of course, I am assuming that foreign speakers are able to make the “ch” sound in German, which some foreigners have difficulty pronouncing.

[16] Another instance of this dogmatic tendency is when, discussing E. O. Wilson’s Sociobiology, he writes, “He [Wilson] is, of course, entitled to define a word however he likes, but this is a most confusing definition, and I hope that Wilson will change it in future editions of his justly influential book.” (94)

[17] If I had to declare what my conception of God is, I would say that it most resembles Albert Einstein’s conception of God, the Creator of the Universe and its many regularities. Although I was raised as a Catholic, I do not believe in any of the central teachings of Christianity.

[18] Although Dawkins would strenuously deny it, I will make the surmise that The Selfish Gene was the result of the superposition of a common science-fiction plot on the Theory of Evolution. If this surmise is correct, it is another similarity between his selfish-gene theory and Sigmund Freud’s theory of psychoanalysis, since the latter was inspired by a perverse misinterpretation of a well-known literary work, Sophocles’ Oedipus Rex.

[19] The Selfish Gene was published in 1976. It would be an unwarranted exaggeration to declare that the views and policies of Margaret Thatcher, who became the prime minister of the UK in 1979, and Ronald Reagan, who became the president of the US in 1981, were influenced by the title of Dawkins’ book. However, I believe that it contributed to the climate pervading the 1980s and subsequent periods, which has been characterized by a greater degree of selfishness and individualism in both the social and economic spheres.

[20] The place where Anjezë Bojaxhiu, as she was then known, was born is now part of Macedonia. After going to Ireland as a teenager to join the Sisters of Loreto, she later went to India, where she founded her own religious order, the Missionaries of Charity.

[21] Although the vast majority of people regard such charitable organisations as entirely good, I do not agree with this view, for they are also largely responsible for the population explosion that is taking place in many poorer countries in the world. There are many undesirable effects of this unchecked human population growth, including war, famine, crime, deforestation, desertification, poaching of endangered species, very high levels of unemployment, and massive urban slums.

[22] In making this statement, I am excepting the behaviours of animals that are determined by imitation, such as in humans and other species, or by other forms of learning. Although the tendency to imitate the behaviours of other members of one’s species is determined genetically, it is obvious that the particular behaviours one observes and imitates are not determined by one’s genes.

[23] In the case of many tamed animals, their tameness is due to early exposure to humans, and not because their usual fear of or hostility towards humans has been bred out. However, in domesticated sheep, both the fight and flight responses have been bred out, as is shown by their inability to survive without human protection, except in places where all large predators have been eliminated by humans.

[24] If some readers find this too great a stretch of the imagination, then one could simply imagine an animal in which the expression of the genes that control for the development of the reproductive organs or mating behaviour is hindered or impaired, something that does occur periodically in the real world. Recently, this is becoming more common due to the chemical and hormonal pollution of the natural world by human beings.

[25] The recent insistence that living organisms should only be considered from the perspective of their genes was probably influenced by the developments in physics during the twentieth century, when the basic components of matter were reduced, from visible substances to atoms, then to protons, neutrons, and electrons, and finally to quarks.

[26] There are many people who go from being atheists or agnostics to believing in God’s existence. These people, who in Christianity are called “born-again Christians,” often seek to convert others to their new-found beliefs. In the case of Richard Dawkins, we have the example of the reverse phenomenon, for he has sought assiduously, with the same moral fervour, to preach his scientific atheism to all non-believers, who in this case are the many who continue to believe in God.

[27] This term has nothing to do with what has been called the “Napoleon complex,” the belief that short persons like Napoleon seek to compensate for their less-than-average height by attempting to accomplish extraordinary feats in order to be noticed by, while distinguishing themselves from, others. This complex is merely a particular instance of Alfred Adler’s theory of the inferiority feeling, or inferiority complex, as it was later popularized.

[28] Because Napoleon had such a high regard for himself, which regard was reinforced by the adulation he received both from his soldiers and, at least for a time, from the French populace, he failed to understand that the other crowned heads of Europe and Russia, although they may have tolerated him and treated him with respect while he was powerful, never regarded him as being their equal, since he had not come to the French throne by – in their eyes – the sole legitimate means of noble birth. This disdain was apparent in the behaviour of his second wife, Marie Louise, the daughter of the Habsburg Emperor Francis II of Austria, who cut all ties with Napoleon and returned to her father with their infant son once it became apparent that Napoleon’s power was steadily and irreversibly declining from its temporary summit.

[29] In my opinion, Sigmund Freud was a literary theorist who masqueraded as a scientist. This would explain why his theory of psychoanalysis has been so fertile and influential in the realms of literary and artistic interpretation and criticism, since it originated from a misinterpretation of a well-known literary work, Sophocles’ Oedipus Rex.

[30] There are many professors, intellectuals, and writers who admire Socrates but fail to realize that if he were alive today, he would go about the corridors of universities, parliaments, news and talk shows, and other places, and confront the most prominent professors, writers, intellectuals, and politicians, and seek to humiliate them by showing that they know nothing. It is much easier to admire someone when one is not subjected to that person’s determined efforts to make one look like a fool, as Socrates did during the latter part of his life.

[31] Although I admit that his theory may have some validity, and that it may help biologists to understand some behaviours, especially non-human behaviours, since, besides humans, many organisms exhibit selfish behaviour, I wholly deny that his theory is valid without any exceptions, which is the position that Dawkins stubbornly maintains.


Introduction

The study of animal behavior has embraced the use of new technologies to understand gene expression and how behavioral phenotypes are produced from the underlying genome (Stapley et al., 2010 Bell and Robinson, 2011). Such approaches will be especially useful for understanding how novel behaviors arise, which has, of course, been a perennial question in evolutionary biology (Darwin, 1859 Mayr, 1982). In some cases the behavior that we observe in animals is not due to the expression of their genes but rather to the genes of parasites infecting them. In such cases the behavior is an extended phenotype of the parasite (Dawkins, 1982 Dawkins, 1990 Dawkins, 2004 Hughes, 2008 Hughes et al., 2012). Beyond the obvious importance of explaining how such complex parasite adaptations evolve by natural selection, the study of behavioral manipulation is important because it represents a parallel experiment over evolutionary time. That is, natural selection has acted on the genome of both the parasite and the host to control a single phenotype (behavior in the host). Understanding diverse pathways from genes to phenotypes will help us tackle the important question in evolutionary biology: what is the mechanistic basis of animal behavior (Duckworth, 2009)? In this Review I explore some of the pathways that can lead us to a proximate level understanding of extended phenotypes.

A crucial detail of extended phenotypes is the distance over which they are extended. This distance can be phylogenetic, as occurs when the parasite and host are distantly related and commonly in different kingdoms [e.g. rabies virus changing mammal behavior (Moore, 2002)]. It is also sometimes a physical distance depending on where in the host’s body the parasite lives [e.g. the abdomen-dwelling hairworms of crickets causing changes in brain expression (Thomas et al., 2002)]. And finally, the distance can be temporal as gene expression of parasite genes may precede the resultant altered phenotype [parasitoids produce chemicals that manipulate insects to act as bodyguards after the wasp has emerged (e.g. Grosman et al., 2008, Maure et al., 2013)]. In spite of these complexities, the task of understanding the genetic basis of an extended phenotype is possible with the correct model system where the biological details are well known (Biron et al., 2005a Lefèvre et al., 2009 Poulin, 2011 Adamo, 2012).

Most studies on parasite manipulation of host behavior have been descriptions of the phenomenon. Unusual behaviors observed in infected individuals are recorded, and if their complexity suggests that it benefits the transmission of parasite genes then the behavior is said to be an example of adaptive manipulation (Barnard and Behnke, 1990 Beckage, 1997 Moore, 2002). This approach is valuable but prone to criticism as adaptationist story-telling (Gould and Lewontin, 1979). Such charges prompted one of the major researchers in the field of parasite ecology to question the utility of the extended phenotype paradigm (Poulin, 2000). In response, over 30 authors debated in a special issue of Behavioral Processes (2005, Vol. 68, Issue 3) that despite problems with adaptationist reasoning, encouragement could be taken from the new studies looking at the mechanisms by which parasites control behavior (Hughes, 2005 Moore et al., 2005 Thomas et al., 2005 Adamo, 2012). This mechanistic approach has already been successful and demonstrated neurological and hormonal changes in infected hosts (Lefèvre et al., 2009 Adamo, 2012). One of the most promising approaches has been an inference of the genetic basis of parasite control via proteomics (Biron et al., 2005a Biron et al., 2005b Biron and Loxdale, 2013). Here, proteome profiles of hairworms (Nematomorpha, also called Gordian worms) causing crickets to jump into water (so that the worms can exit for mating) revealed a molecular cross-talk between the parasite inside the cricket’s abdomen and the brain of the cricket. Specifically, the worms caused an upregulation of cricket Wnt proteins in the brain (Biron et al., 2005b). These advances led to an invited multi-authored review on the mechanistic advances and a call for more detailed studies, including whole-genome analysis (Lefèvre et al., 2009). The field of parasite manipulation has therefore moved beyond its important natural history phase towards a more empirical approach: a recently edited volume for Oxford University Press records these interesting developments and the history leading this point (Hughes et al., 2012).

Although the proteomic basis is important (and the metabolomic basis too), the ultimate goal in studying the extended phenotypes of parasites is to determine the genetic basis. Recently, Hoover et al. (Hoover et al., 2011) were able to demonstrate a single gene effect of baculovirus responsible for an altered behavior (egt), in that case the well-known summit disease observable in virus-infected caterpillars (discussed below). A previous study highlighted the role of a single gene (Cory et al., 2004), but the Hoover et al. study is important for the broader field of parasite manipulation as it points to ways experimental studies (gene knockouts and restoration) will increasingly become part of our toolkit (Fig. 1). Such approaches raise a number of questions for researchers interested in the genetic basis of parasite extended phenotypes that I will discuss in this Review. What I aim to do is ask what different approaches can be taken for better elucidating the genetic evidence for behavioral change. First I will re-cap the concept of the extended phenotype.

The extended phenotype

The paradigm of the gene as the unit of selection emerged during a period of much debate between advocates of individual- and group-level selection and through the work of Bill Hamilton (Hamilton, 1963 Hamilton, 1964). This debate is still on-going and occasionally rancorous (Hughes, 2011). Hamilton’s concepts were subsequently made more transparent by Richard Dawkins in his selfish gene approach (Dawkins, 1976) and became the foundation for sociobiological theory (Wilson, 1975). What this paradigm states is that it is genes alone that are transferred between generations the organisms in which genes reside and their phenotypes are the means by which transmission is secured. Organisms are vehicles and genes are replicators. Natural selection chooses among variation in phenotypes, but the information encoding these phenotypes and, ultimately, the unit that is selected is the gene [see discussion by Mayr (Mayr, 1997)].

The phenotype has principally been considered as a trait of the individual organism. Examples are eye or flower color, antler length, butterfly wing spots, behavior or chemical signals released into the air, to name just a few. But such foci only reflect the convenience with which we could study those easily visible attributes of organisms (Dawkins, 1990). Rapid and continued technological advances allow us to look at phenotypes all the way from the surface of the organism down to the levels of transcription and protein folding. Dawkins (Dawkins, 1982) also advocated an additional level of the phenotype, but what was and still remains novel is that this additional level of phenotype is not physically attached to the organisms whose genes are encoding it this is the extended phenotype.


The genetic diversity of intraterrestrial aliens

In this issue of New Phytologist, Croll et al . (pp. 672–687) describe a study in which they sampled over 40 individuals from a field population and scored each individual for its genotype at each of 13 polymorphic molecular loci. Such studies have been the routine fodder of population genetics since the 1970s, so why is this paper noteworthy? It is because the subjects of this study are arbuscular mycorrhizal fungi (AMF). These are strange organisms, and it has become obligatory to open every paper on AMF with two true statements: AMF are vital for the normal growth of most plants, and AMF are fiendishly difficult to study.

‘AMF are exceptional in that there is no stage in the life history where an individual is reduced to a single nucleus, because the large spores contain hundreds of nuclei.’

The first challenge in AMF population genetics is to sample a set of individuals from the tangled web of root infections in the field, which Croll et al . did by establishing multiple in vitro cultures. Nobody has yet succeeded in growing AMF in pure culture, as the fungus does not grow unless attached to plant roots, but it is possible to propagate AMF on root organ cultures in vitro, although only a few AMF species have so far been grown successfully in this way. This indirect approach is bound to introduce some bias reflecting culturability in the chosen hosts and conditions, but has major advantages. A simpler approach would be to collect spores from the field ( Stukenbrock & Rosendahl, 2005 ), but this is the equivalent of studying a plant community by digging up the seeds in the soil spore numbers are a poor reflection of fungal biomass and activity. The other problem with a spore is that it only has enough DNA for a small number of analyses ( Stukenbrock & Rosendahl, 2005 ), whereas an in vitro culture can be multiplied indefinitely. Croll et al . were therefore able to type each isolate reliably at 13 genetic loci. Furthermore, each locus was targeted by AMF-specific PCR primers, so we can be confident that all the products are from the fungus itself rather than from the bacteria or nonAMF fungi that are frequently closely associated with AMF.

The first surprise is that, at all the loci described by Croll et al ., they report just one allele in each isolate. This might seem a normal expectation for a fungus, given that most fungi are haploid most of the time, but in fact the authors did find some loci that had more than one allele in some individuals, although they did not use them in the study because this would have complicated the analysis (I. Sanders, pers. comm.). The history of multiple sequences within AMF began with the discovery that a single isolate may have two or more distinct sequences for the ribosomal RNA gene region ( Sanders et al., 1995 Lloyd-Macgilp et al., 1996 ). Usually these differences are fairly small, but in the Glomus mosseae group of species, two internal transcribed spacer (ITS) sequences are found, and these are so different that one might expect them to indicate different genera, except that they have been reported to co-occur in the same isolate ( Clapp et al., 2002 ). Nor is within-isolate diversity confined to the ribosomal RNA genes: it has also been seen for other genes ( Helgason et al., 2003 Corradi et al., 2004 ). Whether this diversity reflects the cohabitation of genetically distinct nuclei within the same cytoplasm (a heterokaryon) or identical nuclei, each of which contains all the genetic variants (a homokaryon), is the subject of ongoing debate ( Kuhn et al., 2001 Hijri & Sanders, 2004 Pawlowska & Taylor, 2004 Hijri & Sanders, 2005 Pawlowska, 2005 Corradi et al., 2007 Rosendahl, 2008 ). AMF have few cross-walls within their hyphal networks, so nuclei inhabit a common cytoplasm (a syncytium). This is not so unusual in fungi, but AMF are exceptional in that there is no stage in the life history where an individual is reduced to a single nucleus, because the large spores contain hundreds of nuclei. The nuclei in a spore, and in the mycelium derived from a spore, are therefore better considered as a population rather than as an individual. If they are collectively a homokaryon, we have to explain how the variation generated by mutation is purged, while if they are a heterokaryon, we must explain how diversity is maintained in the face of genetic drift.

There is not space here for a full discussion of all the evidence, so I shall merely point out that the most-cited study supporting the homokaryon hypothesis is far from conclusive. Pawlowska & Taylor (2004 ) studied two genes in Glomus etunicatum. The first, PLS1, was unusual in that 13 allelic variants coexisted in a single isolate. All the variants were maintained through successive generations, which would be unlikely if each variant were in a separate, independently segregating nucleus. However, under some models for heterokaryosis, such as the selfish nucleus hypothesis (Fig. 1), independent segregation of nuclei is not expected. Secondly, Pawlowska & Taylor (2004 ) amplified the ribosomal ITS from a number of individual nuclei, and showed that, whenever they succeeded in getting amplification, a nucleus carried all three of the ITS variants seen in that isolate. Ribosomal genes are unusual in that they are in multiple copies in the genomes of all organisms, and sometimes in several arrays that may diverge in sequence. Admittedly, the sequences in different nuclear lineages within a permanent heterokaryon would be expected to diverge, but Pawlowska & Taylor (2004 ) only succeeded in obtaining amplification for a handful of nuclei, which might have represented a particularly favourable genotype (perhaps with a large number of rDNA copies), rather than a random sample of all the nuclei in the spore. Indeed, in a syncytium, it is theoretically possible for some nuclei to survive even if they have no rDNA at all.

Two hypotheses for genome organization in arbuscular mycorrhizal fungi. (a) The polyploid nucleus hypothesis ( Pawlowska & Taylor, 2004 ) explains how multiple allelic variants can coexist in a homokaryon. Each nucleus has multiple copies of the genome, although massive gene loss frequently follows the formation of a polyploid, so it is likely that individual genes vary widely in copy number. (b) The selfish nucleus hypothesis (proposed here) explains how multiple genetically different nuclei can persist in a heterokaryon without segregational loss. In a syncytium, a nucleus may replicate faster if it loses part of its genome and relies on its neighbours to provide the missing products in the common cytoplasm. The evolutionary result will be a consortium of mutually complementary nuclear lineages, none of which can survive without the others. Other scenarios are also possible for both homokaryons and heterokaryons.

Multiple genetic markers allow a good number of genotypes to be distinguished, and Croll et al . discuss their spatial distribution and association with host species. Furthermore, with multiple loci in multiple individuals, patterns of association between loci can be explored. Here things start to get really interesting. AMF have been described as anciently asexual, because no convincing structures associated with sexual reproduction have been seen, and no unambiguous evidence for genetic recombination ( Rosendahl & Taylor, 1997 ). Ancient asexuals are of interest because most asexual lineages are thought to be short-lived ( Gandolfi et al., 2003 ). This issue, and others raised in this commentary, have been discussed in a recent Tansley review ( Rosendahl, 2008 ). A completely asexual organism evolves as a branching tree of clonal lineages. A genetic variant that arises in one lineage is forever associated with the set of genotypes characteristic of that lineage. Without recombination, it cannot get together with an allele at another locus that arose in a different lineage (Fig. 2). On the face of it, the table of genotypes presented by Croll et al . seems to contradict this expectation. There are several instances where the same allele crops up in genotypes that, on the evidence of the other loci, clearly belong to different lineages. Is this evidence for genetic recombination in the supposedly asexual AMF?

Genetic evidence for asexuality and recombination. In an asexual organism, clonal lineages gradually diverge from each other at all loci. Since all loci share a common ancestry, they all have a consistent phylogeny. The effect of recombination between lineages (indicated by black arrows) is to introduce alleles that have a discordant phylogeny.

The evidence is not conclusive, and the potential problem is homoplasy that is, the possibility that the same allele arose independently more than once. This is a particularly prevalent problem with microsatellites (simple sequence repeats, SSR), and most of the loci in the study were of this kind. An SSR locus consists of a number of tandem repeats of a very short DNA motif, typically two or three bases. Mutations that increase or decrease the number of repeats occur frequently by strand-slippage during replication, so such loci are typically highly polymorphic in populations. This makes them handy as genetic markers, but homoplasy is rife at SSR loci because a variant with a particular number of repeats can be generated in many ways. Hence, SSRs do not seem a good choice for a critical study of interlocus associations. However, an examination of the allele sequences that Croll et al . provide in their supplementary material reveals that, although they chose loci because they included SSR motifs, most of the allelic differences that they observed are not the result of typical SSR length variation. In fact, the alleles differ in multiple ways, including short insertions or deletions and single nucleotide substitutions. When an allele has multiple unique features, it is very unlikely to have arisen more than once, so we can discount homoplasy. Some ambiguity remains, though, because Croll et al . do not provide a complete sequence for every gene in every individual. Most individuals are only characterized by the overall length of the PCR product from each locus. It is easy to see that quite different sequences can happen to be of the same overall length and, in the case of the nuclear intron locus, the authors demonstrate exactly this. Nevertheless, Croll et al . appear to have the tools for a rigorous assessment of recombination in AMF. Will we soon have to reassess the assumption that AMF are ancient asexuals?


On the Evolution of Human Morality

Steven Pinker’s thoughtful remarks concerning group selection present a useful occasion for clearing some misconceptions surrounding recent developments in the behavioral sciences concerning our understanding of moral vs. self-interested behavior. Initiated in 1966 by George C. Willams’ Adaptation and Natural Selection and followed a decade later by Richard Dawkins’ The Selfish Gene, evolutionary biologists in the last quarter of the Twentieth century came to view humans as fundamentally selfish, contributing to society only when socially-imposed rewards and punishment render it in their self-interest to do so. Dawkins, for instance, opines in the opening pages of The Selfish Gene, “We are survival machines—robot vehicles blindly programmed to preserve the selfish molecules known as genes…. a predominant quality to be expected in a successful gene is ruthless selfishness. This gene selfishness will usually give rise to selfishness in individual behavior…. Anything that has evolved by natural selection should be selfish.”

Of course, it does not appear in our daily life that everyone is selfish, and if we introspect, most of us will agree that we try to behave, however successfully or unsuccessfully, as moral beings willing to sacrifice personal amenities in the pursuit of truth, justice, loyalty and compassion. Dawkins’ explanation is that human morality is a cultural facade laid upon our basically selfish human nature. “Be warned,” he states, “that if you wish, as I do, to build a society in which individuals cooperate generously and unselfishly towards a common good, you can expect little help from biological nature. Let us try to teach generosity and altruism, because we are born selfish.”

But why do fundamentally selfish beings, which is what humans are according to the selfish gene theory, accept cultural norms that contradict their natural strivings? Richard Alexander answered this question in 1987 in his The Biology of Moral Systems with his concept of indirect reciprocity, according to which we all continually evaluate others for possible future gainful interactions, and we reject individuals who violate norms of reciprocity. The somewhat more general answer offered by Pinker is that is that each of us conforms to social norms out of fear of losing our good reputation. What appears to be self-sacrifice is thus simply a superficial veneer covering our selfish natures. “Scratch an altruist,” biologist Michael Ghislin eloquently wrote in 1974, “and watch a hypocrite bleed.”

Pinker frames the issue in terms of sacrificing personal interests on behalf of the group. “What we don’t expect to see,” he writes, “is the evolution of an innate tendency among individuals to predictably sacrifice their expected interests for the interests of the group.” This is not the correct way to frame the issue. People do not generally “sacrifice on behalf of the group.” Rather, people have moral principles that the strive to uphold, and that compete with their material interests. When I behave honestly in a transaction, I may have no intention whatsoever of sacrificing on behalf of my transaction partners, much less on behalf of my society. I just do what I think is the morally correct thing to do. When I bravely participate in a collective action against a despotic regime, I am upholding my moral principles, not sacrificing on behalf of the group. Indeed, it is no sacrifice at all to behave morally, because we humans care about our moral worth in much the same way as we care about our material circumstances.

The past few decades have seen the massive accumulation of evidence in favor of the view that human beings are inherently moral creatures, and that morality is not a simple cultural veneer. Humans are born with a moral sense as well with a predisposition to accept and internalize moral norms their society, and often to act on these moral precepts at personal cost. In our book, A Cooperative Species, Samuel Bowles and I summarize a plausible model of human nature in which “people think that cooperating is the right thing to do and enjoy doing it, and that they dislike unfair treatment and enjoy punishing those who violate norms of fairness.” Most individuals include moral as well as material goals in their personal decision-making, and they willingly sacrifice material interests towards attaining moral goals. It is this view that I will defend in my remarks.

Pinker does not present, and indeed makes light of the body of research supporting the existence of a basic human moral sense, suggesting that there is only one piece of evidence supporting the view that people behave morally when their reputations are not at stake: “It seems hard to believe,” he says, “that a small effect in one condition of a somewhat contrived psychology experiment would be sufficient reason to revise the modern theory of evolution, and indeed there is no reason to believe it. Subsequent experiments have shown that most of the behavior in these and similar games can be explained by an expectation of reciprocity or a concern with reputation.” Because expectation of reciprocity and concern for reputation are basically selfish and do not involve a fundamental respect for moral values, Pinker is simply reiterating Dawkins’ message of a half-century ago that we are the selfish product of selfish genes.

1. Morality and Human Nature
Today the economics and psychology journals, including the most influential natural science journals, Science and Nature, are full of accounts of human moral and prosocial behavior. Pinker dismisses this evidence by asserting that “Any residue of pure altruism” beyond self-interested reciprocity and reputation building “can be explained by the assumption that people’s cooperative intuitions have been shaped in a world in which neither anonymity nor one-shot encounters can be guaranteed.” In other words what looks like moral behavior is just a form of mental error due to imperfections of the human brain.

The empirical evidence on cooperation in humans does not support Pinker’s view. The social interactions studied in the laboratory and field always involve anonymity, so subjects cannot help or harm their reputations, and they usually are one-shot, meaning that subjects cannot expect to be rewarded in the future for sacrifices they make at a given point in time.

Pinker does cite a few studies that support his position. “Subsequent experiments have shown that most of the behavior in these and similar games can be explained by an expectation of reciprocity or a concern with reputation.” Let us see what these studies in fact say. Reciprocity, says Pinker, “is driven not by infallible knowledge but by probabilistic cues. This means that people may extend favors to other people with whom they will never in fact interact with again, as long as the situation is representative of ones in which they may interact with them again.” The only published paper he cites is by Andrew W. Delton, Max M. Krasnow, Leda Cosmides and John Tooby, “Evolution of Direct Reciprocity Under Uncertainty can Explain Human Generosity in One-shot Encounters.”[1] This paper (and several related papers coming out of the Center for Evolutionary Psychology at Santa Barbara, California) show that, in the authors’ words “generosity is the necessary byproduct of selection on decision systems for regulating dyadic reciprocity under conditions of uncertainty. In deciding whether to engage in dyadic reciprocity, these systems must balance (i) the costs of mistaking a one-shot interaction for a repeated interaction (hence, risking a single chance of being exploited) with (ii) the far greater costs of mistaking a repeated interaction for a one-shot interaction (thereby precluding benefits from multiple future cooperative interactions). This asymmetry builds organisms naturally selected to cooperate even when exposed to cues that they are in oneshot interactions.”

This statement is of course not only true, but completely obvious, and does not require sophisticated academic papers to validate its truth. However it does not explains human generosity. It is elementary logic that to say that P explains Q does not mean that if P is true then Q is true, but rather the converse: whenever Q is true, then P is true as well. In the current context, this means that whenever subject A sacrifices on behalf of stranger B in an experiment, it must be true that A is sufficiently uncertain concerning the probability of meeting B again, and A would incur a sufficiently large cost should A meet B again in the future, that it pays A to sacrifice now. The authors have not even attempted to show that this is the case. Nor is it plausible. The experiments under discussion assume subject anonymity, subjects will never knowingly meet again. Pinker’s supposed counter-evidence is thus invalid. To my knowledge, there is simply no credible counter-evidence.

2. Human Morality in Everyday Life
Many readers will doubtless wonder if our view of human moral behavior, which is based on controlled laboratory and field studies, extends to real life. Consider, for one example among many, political activity in modern societies.

In large democratic elections, the selfish individual will not vote because the costs of voting are positive and significant, but the probability that one vote will alter the outcome of the election is vanishingly small. Thus the personal gain from voting is vanishingly small. The cost, however, is a significant amount of time and energy that could have been devoted to other, materially rewarding, purposes. It follows also that a selfish individual will generally not bother to form opinions on political issues, because these opinions cannot affect the outcome of elections.

Yet people do vote, and many do expend time and energy in forming political opinions. This behavior does not conform to the selfish gene model. Of course it could be argued that we only vote to enhance our reputation as a good citizen, but since who votes is normally not public information, and one’s voting history of little interest to employers and other social intimates, this is not a very plausible explanation.

It is a short step from the irrefutable logic of selfish political behavior that selfish individuals will not participate in the sort of collective actions that are responsible for the growth of representative and democratic governance, the respect for civil liberties, the rights of minorities and women in public life, and the like, that are characteristic of many modern societies. In the selfish gene model, only small groups of individuals who seek social dominance will act politically. Yet modern egalitarian political institutions are the result of such collective actions. This behavior cannot be explained by the selfish gene model.

Except for professional politicians and socially influential individuals, electoral politics is a vast morality play to which models of the selfish actor are a very poor fit.

Defenders of the selfish gene theory may respond that voters believe their votes make a difference, however untenable this belief might be under logical scrutiny. Indeed, when asked why they vote, voters’ common response is that they are trying to help get one or another party elected to office. When reminded that one vote cannot make a difference, the common reply is that there are in fact close elections, where the balance is tipped in one direction or another by only a few hundred votes. When reminded that one vote will not affect even such close elections, the common repost is that “Well, if everyone thought like that, we couldn’t run a democracy.” Agreed. But this is just the Kantian categorical imperative, an eminently moral value. People vote because it is simply the right thing to do.

Politically active and informed citizens appear to operate on the principle that voting is both a duty and prerogative of citizenship, an altruistic act that is justified by the categorical imperative: act in conformance with the morally correct behavior for individuals in one’s position, without regard to personal costs and benefits. Such mental reasoning, which has been called “shared intentionality,” is implicated in many uniquely human cognitive characteristics, including cumulative culture and language. Shared intentionality rests on a fundamentally prosocial disposition.

Human beings acting in the public sphere are, then, neither avid reputation mongers nor personal gain maximizers. Rather, they are in general what Aristotle called zoon politikon—political beings. And political beings are moral beings.

3. Cultural Evolution Theory is not Just History
“Most of the groupwide traits that group selectionists try to explain,” says Pinker, “are cultural rather than genetic…. Instead, they are traits that are propagated culturally… group selection … is not a precise implementation of the theory of natural selection… Instead it is a loose metaphor, more like the struggle among kinds of tires or telephones. For this reason the term `group selection’ adds little to what we have always called `history’.” There are two misconceptions in this statement. The first is that group selectionists are for the most part uninterested in genetic evolution. The second is that the concept of cultural evolution is a simply a metaphor.

In fact, the general framework within which we work is called gene-culture coevolution. This framework is quite clearly delineated in scientific terms (see, for instance, my overview, “Gene-culture Coevolution and the Nature of Human Sociality”[2] which I can summarize as follows.

First, as elucidated by Richard Lewontin in 1970, natural selection applies to any entity that follows certain rules. “The principle of natural selection as the motive force for evolution… embodies three principles.” These principles are first, phenotypic variation: “Different individuals in a popUlation have different morphologies, physiologies, and behaviors.” Second, differential fitness: “Different phenotypes have different rates of survival and reproduction in different environments.” And finally, fitness must be heritable: “There is a correlation between parents and offspring in the contribution of each to future generations.” Note that there is nothing about genes in this account of natural selection. Indeed, Darwin himself never heard of genes when he wrote The Origin of Species by Means of Natural Selection. “It is important to note,” writes Lewontin, “a certain generality in the principles. No particular mechanism of inheritance is specified… The population would evolve whether the correlation between parent and offspring arose from Mendelian, cytoplasmic, or cultural inheritance.”

Second, many forms of culture do indeed follow these three principles. Different societies have different cultural norms and technologies (phenotypic variation), some cultural forms contribute more to the fitness of individuals in the societies that embrace these forms, and therefore the cultural forms themselves have varying rates of having copies in succeeding generations (differential fitness), and finally, there is a degree of faithfulness in copying cultural objects from one generation to the next (heritability).

Third, there is a common underlying unity to genetic and cultural evolution. Genes transmit information from one generation to the next, encoded in DNA, that is used by new individuals for ontological development and adaptation to the environment. Cultural forms transmit information from one generation to the next, encoded in human brains, artifacts, and documents, used by new individuals for similar purposes.

Finally, when humans develop new cultural forms, such as language, tools, lethal weapons, control of fire and cooking of edibles, the long-term effect is the transformation of the human genome itself. In other words, cultural evolution leads to genetic evolution. Examples are cooking, which led to a vast reduction in the size of the human gut, language, which led to radical changes in the human larynx and tongue as well as a considerable increase in brain size, and lethal projectile weapons, which led to changes in the morphology of hand and shoulders, as well as a reorganization of the upper torso musculature. Consider, for instance, that our closest relative, the chimpanzee, spends five hours a day digesting comparing to one for humans, lacks the ability to produce complex vocalizations, and cannot throw a stone with more than minimal accuracy or force.

Gene-culture coevolution provides a plausible scenario for the development a moral sense in humans, a quality that appears to be absent or extremely rudimentary in other species. Of course, we may never know with certainty because the paleoanthropological record is extremely scanty. However, it is clear that the development of tools, weapons, and cultural objects required cultural norms promoting cooperation. Individuals who violated these norms were doubtless punished and shunned, and hence less likely to pass their genes on to the next generation. A genetic predisposition to conform to social norms was thus likely to be biologically fit, and hence to evolve through natural selection. Insofar as social norms contribute to the fitness of the groups that embrace these norms, so will the genetic predisposition to follow these norms. We call such a predisposition a moral sense.

4. The Group Selection Pseudo-controversy
Pinker begins his discussion of the group selection issue with the following question. “Human beings live in groups, are affected by the fortunes of their groups, and sometimes make sacrifices that benefit their groups. Does this mean that the human brain has been shaped by natural selection to promote the welfare of the group in competition with other groups, even when it damages the welfare of the person and his or her kin? If so, does the theory of natural selection have to be revamped to designate `groups’ as units of selection, analogous to the role played in the theory by genes?” There are two misconceptions in the very posing of this question.

If an altruistic behavior reduces the net fitness of the altruist and his kin, it cannot evolve. The “group selectionists” argue that while the altruist may be less fit that the selfish individuals in his group, groups with many altruists will expand at the expense of groups with few or no altruists, and this expansion can more than offset the fitness loss of the altruist. Because the altruists kin are more likely to be altruists and also are more likely to be in the altruist’s (this is called `limited dispersal’), the net effect of the altruistic act may be to increase the average fitness of the altruist’s kin.

The first misconception here is the view that group selection is incompatible with kin selection. It is not. Kin selection says that the fitness of an individual depends on the genes of his kin and not just his own genes. Group selection says the fitness of an individual depends on the characteristics of the group he is in, not just his own genes. The second misconception is that group selection means that the group is a “unit of selection.” This is not true. Group selection occurs when the fitness of individuals may be higher in one group rather than other, depending on the social structure of the group and its and distribution of genomes.

A third misconception is that if genes are the only true replicators in evolutionary biology, and if genes are in some sense purely selfish replicators, then all biological species must ultimately sacrifice only for their close genealogical relatives. We can thus admit the complex division of labor and altruism in such eusocial species as termites and honeybees, but we must deny altruism in the case of humans, who cooperate widely with non-relatives. In fact, a careful development of gene-level fitness dynamics in a recent paper by Andy Gardner and J. J. Welsh, “A Formal Theory of the Selfish Genootnote Journal of Evolutionary Biology, 24, 2011. shows that even an inclusive fitness maximizing selfish gene can support altruistic behavior in its owner.

5. The Group Selection Issue is Scientific, not Political
The group selectionists, says Pinker “have drawn normative moral and political conclusions from these scientific beliefs, such as that we should recognize the wisdom behind conservative values, like religiosity, patriotism, and puritanism, and that we should valorize a communitarian loyalty and sacrifice for the good of the group over an every-man-for-himself individualism.” If this were true, it would, in my eyes, be a violation of scientific principles. Even if every known society has followed a certain practice (for instance, eating meat), and even if we can find evolutionary roots for this practice, it would not follow that this practice is morally defensible or practically desirable in our society. Indeed,, there is not a single paper or book that I authored or coauthored that drew any such political conclusions, or indeed any political conclusions at all. Nor am I aware of others who work in this tradition who have drawn such conclusions. Sociobiology is not liberal or conservative, or even middle of the road. It is just good science.

6. Conclusion
Some academic fields, including sociology, anthropology, and psychology, have no problem dealing with the fact that human beings are moral creatures, and that this morality is an important element in our success as a species. Others, including evolutionary and population biology, have had a harder time with this fact, because they have over the years developed theories that appear to show that evolutionarily fit behavior is necessarily selfish behavior. However, the moral nature of human society, and the key role of morality in our success as a species, can be accommodated without requiring evolutionary biology to abandon its cherished accomplishments. The selfish gene versus group selection issue, when properly formulated, has little to do with the nature of human sociality.

[1] Proceedings of the National Academy of Sciences 108,32 2011.


Richard Dawkins

Richard Dawkins is Professor of the Public Understanding of Science at Oxford University. He is the author of many books including the international best-sellers " The Selfish Gene", "The Blind Watchmaker", and "Climbing Mount Improbable."

QUESTION: Professor Dawkins, could you explain your belief that human beings are just "gene machines"?

MR. DAWKINS: When I say that human beings are just gene machines, one shouldn't put too much emphasis on the word "just." There is a very great deal of complication, and indeed beauty in being a gene machine. What it means is that natural selection, Darwinian natural selection, which is the process that has brought all living things to be the way they are, is best seen at the gene level, is best seen as a process of differential survival among genes, and therefore living organisms and their bodies are best seen as machines programmed by the genes to propagate those very same genes. In that sense we are gene machines. But it is not intended to be at all a demeaning or belittling statement.

QUESTION: Now, if we are gene machines, presumably then our behavior is also programmed by genes -- you have made that case. But Christians would say that there is a thing called free will, and that free will gives us a genuine choice about our actions, that effectively free will allows us to override biology. What is your response to that as a scientist?

MR. DAWKINS: I am very comfortable with the idea that we can override biology with free will. Indeed, I encourage people all the time to do it. Much of the message of my first book, "The Selfish Gene," was that we must understand what it means to be a gene machine, what it means to be programmed by genes, so that we are better equipped to escape, so that we are better equipped to use our big brains, use our conscience intelligence, to depart from the dictates of the selfish genes and to build for ourselves a new kind of life which as far as I am concerned the more un-Darwinian it is the better, because the Darwinian world in which our ancestors were selected is a very unpleasant world. Nature really is red in tooth and claw. And when we sit down together to argue out and discuss and decide upon how we want to run our societies, I think we should hold up Darwinism as an awful warning for how we should not organize our societies.

QUESTION: So you are not saying then that our genetic programming is fully deterministic?

MR. DAWKINS: It's an important point to realize that the genetic programming of our lives is not fully deterministic. It is statistical -- it is in any animal merely statistical -- not deterministic. Even if you are in some sense a determinist -- and philosophically speaking many of us may be -- that doesn't mean we have to behave as if we are determinists, because the world is so complicated, and especially human brains are so complicated, that we behave as if we are not deterministic, and we feel as if we are not deterministic -- and that's all that matters. In any case, adding the word "genetic" to deterministic doesn't make it any more deterministic. If you are a philosophical determinist, then adding the word "gene" doesn't increase the effect.

QUESTION: If, as you have said, there is a tendency from our genes for us to be selfish, would that perhaps suggest that we need institutions like religion to encourage us to override this innate selfishness?

MR. DAWKINS: The phrase "the selfish gene" only means that genes are selfish. It doesn't mean that individual organisms are. On the contrary, one of the main messages of the selfish gene is that selfish genes can program altruistic behavior in organisms. Organisms can behave altruistically towards other organisms -- the better to forward the propagation of their own selfish genes. What you cannot have is a gene that sacrifices itself for the benefit of other genes. What you can have is a gene that makes organisms sacrifice themselves for other organisms under the influence of selfish genes.

I think we certainly benefit from social institutions which encourage us towards moral behavior. It's very important to have law. It's very important to have a moral education. It's very important to try to inculcate into children moral rules, such as "do as you would be done by." It's very important to do moral philosophy, to try work out the principles we want to live. But when you say religious principles, there I think I would part company. I see no reason why they should be religious. But I certainly think that they should be developed by society and not necessarily following biological dictates.

QUESTION: What is your response to the view that some Christians are putting forward that God is the designer of the whole evolutionary system itself?

MR. DAWKINS: In the 19th century people disagreed with the principle of evolution, because it seemed to undermine their faith in God. Now there is a new way of trying to reinstate God, which is to say, well, we can see that evolution is true. Anybody who is not ignorant or a fool can see that evolution is true. So we smuggle God back in by suggesting that he set up the conditions in which evolution might take place. I find this a rather pathetic argument. For one thing, if I were God wanting to make a human being, I would do it by a more direct way rather than by evolution. Why deliberately set it up in the one way which makes it look as though you don't exist? It seems remarkably roundabout not to say a deceptive way of doing things.

But the other point is it's a superfluous part of the explanation. The whole point -- the whole beauty of the Darwinian explanation for life is that it's self-sufficient. You start with essentially nothing -- you start with something very, very simple -- the origin of the Earth. And from that, by slow gradual degrees, as I put it "climbing mount improbable" -- by slow gradual degree you build up from simple beginnings and simple needs easy to understand, up to complicated endings like ourselves and kangaroos.

Now, the beauty of that is that it works. Every stage is explained, every stage is understood. Nothing extra, nothing extraneous needs to be smuggled in. It all works and it all -- it's a satisfying explanation. Now, smuggling in a God who sets it all up in the first place, or who supervises the details, is simply to smuggle in an entity of the very kind that we are trying to explain -- namely, a complicated and beautifully designed higher intelligence. That's what we are trying to explain. We have a good explanation. Why smuggle in a superfluous adjunct which is unnecessary? It doesn't add anything to the explanation.

QUESTION: What do you say to the argument that some people are raising now that it's all very well for evolution to be the mechanism once you have a self-replicating structure like DNA -- but how do you get that complex structure in the first place? Maybe DNA is the work of God?

MR. DAWKINS: It's a different argument to say how did the whole process start - how do we begin with the origin of life? The origin of life -- the key process in the origin of life was the arising of a self-replicating molecule. This was a very simple thing compared with what it's given rise to. By far the majority of the work in producing the elegant complexity of life is done after the origin of life, during the process of evolution. There does remain the very first step -- I don't think it's necessarily a bigger step than several of the subsequent steps, but it is a step. And it's a step which we don't fully understand -- mainly because it happened such a long time ago, and under conditions when the Earth was very different. And so it's not necessarily possible to simulate again the chemical conditions of the origin of life. There are various theories for how it might have happened. None of them is yet fully convincing. It may be that none of them ever will be, because it may be that we shall never know fully what the conditions were. But I don’t find it at all a deeply mysterious step.

QUESTION: You have called religion a virus. What do you mean by that?

MR. DAWKINS: The word "virus" is used in various ways. The original way is the proper way like the influenza virus or the measles virus -- a bit of DNA which gets into the cell and programs it to its own end. Computer viruses are an analogy -- it's a very good analogy, because a computer virus is a piece of computer code written in computer language. It says, "Duplicate me and spread me around and maybe do some mischief on the way," and it works because computers obey the instructions written in computer language. If you write a program that says "duplicate me, spread me around," it will spread by the medium of floppy disks and so on.

Now, the human brain may be vulnerable to being parasitized in the same kind of way as the cell is by DNA viruses, and as computers are by computer viruses. An example would be the chain letter -- the letter that you get through the post and says, "Make 10 copies of me, and send me to 10 friends." And of course if everybody literally obeyed slavishly, like a computer would, then the world would soon be knee-deep in paper, because it's a very rapid exponential increase in bits of paper. Humans aren't quite that stupid. But they are sufficiently stupid to fall for certain kinds of inducements, such as, "If you don't believe this and pass it along you'll go to hell when you die." That's a threat. It's a threat that sufficient numbers of people believe to agree to pass on the equivalent of the chain letter --the message, the gospel, whatever it may be.

There's an added inducement, an added fact, which is that the brains of children, I think for probably very good Darwinian reasons, are especially susceptible for good reasons. A child, a young child, needs to learn a great deal -- not only a language, but lots of other things, from its culture -- from the elders of its society, from its parents. This means that children are going to be pre-programmed to believe whatever they're told at a certain tender, vulnerable age. And I use the word "vulnerable," because automatically like the computer they are then going to be vulnerable to parasitization by any other kind of information that is given them, which may not be good information. So the information, "Don't put your hand in the fire, because you'll be burned," or "Don't tread on yellow-and-black-striped snakes, because you'll be bitten" -- that's good information.

The child has no way of knowing what's good information. And so some other piece of information "X" -- it doesn't matter what it is -- it may be completely nonsensical. But if the child is told at a certain tender age, that when you grow up you must pass on the same X to your own children, then that X is going to get passed on from generation to generation, regardless of what it is. And you predict from this that what people believe in one part of the world will be arbitrarily different from what people believe in another part of the world -- nothing to do with difference in truth. There is no difference in the truth about the cosmos in Roman Catholic countries or Protestant countries or Islamic countries. Yet children in Islamic countries are told one thing and children in Catholic countries are told another thing, and they all believe what they are told. At least for a while some of them manage to shake it off later -- but many don't so the message gets passed down the generations indefinitely -- and I think that's one of the things that explain the existence of religion.

QUESTION: What is your view of more liberal religious views that are held by people like your Oxford colleage Arthur Peacocke, who is both a biochemist and an Anglican minister?

MR. DAWKINS: More sophisticated theological views, people like Arthur Peacocke and John Polkinghorne -- obviously they're not creationists in any simple sense -- they're not fundamentalists, they're not stupid. So do I respect them more? Well, in one respect obviously I do, because really you could have an intelligent conversation with them -- they're not ignorant. On the other hand, I can't understand what they're doing it for. I mean, I don't understand what it is that is being added, either to their lives or to the storehouse of human wisdom by bringing in this additional dimension of explanation. We have science. Science is by no means complete -- there's a lot that we don't know -- but we're working on it. Both of those two gentlemen are scientists, and they know what that means. They understand it and they respect it. We're working on building up a complete picture of the universe, which if we succeed will be a complete understanding of the universe and everything that is in it. So I don't understand why they waste their time going into this other stuff which never has added anything to the storehouse of human wisdom, and I don't see that it ever will.

QUESTION: If you could wave a magic wand and have the world be the way you think it should be, what would you see as being the proper relationship between science and religion today?

MR. DAWKINS: If I am asked is there a role for religion, would I just like to wave a magic wand and wipe it out? It's not quite so straightforward as that. It could sound patronizing, but I could imagine that the consequences of simply wiping it out with a magic wand could -- there could be social, psychological consequences which could be rather drastic. I mean, there are people who have come to depend upon it. There are people who rely upon it like a crutch. And so simply to wipe it out might be rather like taking the placebo pills that somebody has been used to taking all their lives and which have kept them happy and healthy, and suddenly saying, "Sorry, you can't take your pills anymore." So I wouldn't wish to be so glib as to say I just want to wipe it out.

But as far as serious intellectual discourse is concerned, as far as actually trying to understand the world where humanity is concerned, yes, I think we are wasting our time doing that. I don't mean to say that all theologians are wasting their time -- I should have said that earlier. I mean, it's important to recognize that what theologians actually do in many cases is not what I call theology -- it's Biblical history or literary criticism of ancient Hebrew text, and that's fascinating and should be done, just as the same exercise on English historical texts should be done. I'm all for that. I love history, I love literature, and I love Biblical history and Biblical literature, and I am very keen that that should be done as a proper branch of scholarship.

But theology as opposed to Biblical history and literature -- when you argue about the true inner meaning of the trinity, or the transubstantiation, and try to come up with some symbolic meaning -- I think that is a total waste of time. I mean, a good satirical parallel might be to suppose that one day in the fullness of time science discovers that the DNA double helix is false, that we got it all wrong, and DNA is not a double helix. Now, any scientist would say, "Right, pity about that, but we'll now work on finding out what it really is." My satire on theology would be: "Ah, but in some other sense the DNA double helix surely has some meaning for us. What is the DNA double helix trying to tell us in the world today?" Maybe the twisting of the two strands of DNA has some significance for the uniting of human beings one with another -- we must set aside the purely mundane issue of is it true, which is crude and facile -- we are not talking about truth in any simple sense -- we want to find the underlying symbolic truth. There never was an underlying symbolic truth. Either it's true or it isn't.

At the present we think DNA really is a double helix. If ever that's found to be false we throw it out of the window and we start again, and we don't try to rediscover some inner symbolic meaning, which is exactly what they're trying to do with things like the Book of Genesis. They have thrown it out as historical fact, which is what it always was thought to be, and which many of its authors presumably intended it to be -- and they have now replaced it with a symbolic meaning: the true meaning of the Book of Genesis is this that or the other. You know the kind of thing I'm talking about. I think that it is a waste of time. I think it's nonsense.

QUESTION: Why do you think that in an age of science so many people, even in the West, and particularly in America, continue to believe in religion?

MR. DAWKINS: I don't understand why so many people who are sophisticated in science go on believing in God. I wish I did. You'd have to ask them. I know that in some cases what they mean by God is very different from what the ordinary people that they talk to think they mean by God. There are physicists who are deeply awed, as I am, by the majesty of the universe, by the mystery of origins -- the origins of the laws of physics, the fundamental constants of physics, and who are moved by this to say there is something so mysterious that it is almost like God, and maybe use the metaphor of God. God is in the equations. God is in the fundamental constants. And that's fine. I mean, that's just redefinition of that which we find mysterious at the basis of the universe.

But other people misunderstand that, because to them God is that which forgives sins, that which transubstantiates wine, that which makes me live after I died -- and that is a totally different matter. And yet the misunderstanding is ripe for the picking. People will listen to sophisticated physicists, using God as this kind of metaphor for the deep constants, the deep problems, the deep principles of physics, and say that in that sense I believe in God. The reaction is, "Oh, this great physicist believes in God -- that means I'm free to believe in the trinity and in the crucifixion and in the reincarnation of Christ -- and all that stuff, which of course has nothing whatever to do with the fundamental constants of physics, which is what these physicists are talking about.

QUESTION: I want to come back to the issue of viruses, because a theologian once said to me, "If religion is a virus, then we must accept the concept that science might also be a virus." What do you say to that?

MR. DAWKINS: It's often said that if we say that religion is a virus, then why isn't science a virus? Coming back to the metaphor of computer viruses, you could say that any computer program is a virus, even a really good word processor that everybody buys because they need it -- is a virus. It spreads because everybody wants it. The difference is that a computer virus spreads not because everybody wants it, but because it simply says in its instructions, "Spread me." It's a difference of simplicity and pointlessness. The word processor program spreads because it's a good program, because it's sophisticated, because people find that it works, finds that it improves their productivity -- all sorts of complicated reasons why it spreads.

Now, that's analogous to why some elephants survive better than other elephants or some kangaroos survive better than other kangaroos. Fundamentally at the gene level elephants are just like gigantic colonies of viruses which are all programming the elephant to make more DNA via the indirect process of making an elephant first. But it doesn't do justice to the process, because an elephant is such a complicated creature, such a complicated thing, that we focus our attention on the elephant, and we see that it is doing it by another means. And the reason why one elephant survives better than another is it has a superior trunk or better feeding behavior, or sharper tusks. There are all sorts of complicated and elegantly beautiful reasons why one elephant passes on its DNA better than another elephant. But with a virus - a measles virus or a common cold virus, it is not like that. It is exceedingly simple. The virus just says, "Make copies of me." "Make copies of me." And so the machinery obeys it.

Now, science it seems to me is like the elegant word processor. A scientific idea -- some die, some live -- there is a kind of Darwinism of scientific ideas. Philosophers such as Pappas have made the point. But the reason why they spread, the reason why some die and some live, is that they are tested, they are good ideas, they inspire people. They survive the test of experiment. And models built from them lead to predictions which are tested and so on -- a complicated, sophisticated business, like an elephant. Whereas religious ideas, such as "If you don't believe in this you'll go to hell," seem to me to be crudely self-serving in the same kind of way as a computer virus is. I suppose you could make the case that a whole church, the whole Russian Orthodox Church, or the whole Roman Catholic Church, is a sufficiently complicated colony of such viruses that it starts to look a bit more like an elephant, and maybe that's why they survive so well. And I'm happy to go along with that analogy. But I don't think it's helpful to use a demeaning word like "virus" for an elephant, and I don't think it's helpful to use a demeaning word like "virus" for science.

QUESTION: Going back to the notion of genes - to what extent do you think human behavior is geneticly determined?

MR. DAWKINS: Much of my work has been on animal behavior and much of my writing has been on animal behavior, and so although I think of the whole organism as a gene machine, I particularly concentrate on its behavior as one of the main, most interesting manifestations. And so much of my writings are about why this behavior or that behavior in the animal furthers the interests of the genes that built the nervous system that makes it behave in that way.

Now, the question arises: Do we do the same thing for humans, or should we do the same thing for humans? Well, humans are animals -- we're apes -- we're African apes. On the face of it there is no reason why we shouldn't. But for any animal we have to ask that kind of question in a sophisticated way. For one thing, any animal may now be living in a different environment from the environment in which its ancestors lived and in which its genes are naturally selected. And so what you may be seeing is behavior which if you interpret it correctly would have been doing something good for the genes back in the natural environment, but which is now doing something else which is only good for the genes in a sort of indirect, sophisticated sense.

With humans there's no problem in seeing why sexual desire has Darwinian survival value. Anybody can see that. And we therefore have little problem in generalizing it too. We no longer go around with a club and sort of beat the female over the head with it, or whatever our ancestors did. But we now use sophisticated modern means of achieving the same end. It doesn't mean that there are genes for buying smart sports cars the better to impress a member of the opposite sex. There are genes for intent -- for doing whatever is necessary to impress the opposite sex, in whatever environment you find yourself in. And since the environment changes the means will change. Now, that's still a relatively simple, rather crude example -- may even given offense to people, and I don't mind in the least.

But there will be other kinds of things which are less obvious -- like people may think about why business executives are said to demand bigger and bigger desks as they get more senior, or bigger carpets, or whatever it may be. And you can come up with amusing explanations like, the bigger desk in a modern-day environment would be equivalent to something else in the Pleistocene of Africa when the natural selection was going on. And now it's interpreted as a bigger desk, because we have big brains and we live in a world of desks and office blocks and telephones and things, which are quite different from those in which our genes were naturally selected. But you mustn't attempt to give a Darwinian interpretation to every detail.

Moreover, this is now a quite different point -- when brains became sufficiently big they took off in other directions, which no longer have really any connection with gene survival at all. We drive ourselves to achieve goals -- like painting a picture or writing a book, which by almost any stretch of the imagination cannot be said to directly benefit our survival or that of our genes. Perhaps you could say the tendency to drive oneself towards a goal has Darwinian survival value. The tendency to set up a target which one then works by all means available to try to achieve -- perhaps that has Darwinian survival value. But nowadays the targets we set up are things like finishing a book, whereas the target setting-up engine, which is in our brain, was originally set up for a different purpose, like catching a buffalo or finding a water hole. So the mechanism was put there for one purpose, but now our brains have gotten sufficiently big that it has taken off into other directions. And this really gets me back to what I was saying before, about free will and the fact that we can deliberately take the decision to emancipate ourselves from the world of natural selection in which genes were naturally selected, and make a new world for ourselves to live in, which is an anti-Darwinian world.

QUESTION: In what way could we liberate ourselves from the Darwinian world? If this is a philosophical issue as opposed to a scientific one, what do you mean by that?

MR. DAWKINS: Some people are puzzled by the sense in which it's possible to take a deliberate decision to emancipate ourselves from the Darwinian world. Well, we know we do it, because every time we look unreceptive we are doing something anti-Darwinian. What happens is that Darwinian natural selection has built into us sexual desire for obvious good Darwinian reasons. In nature, where there are no contraceptives, sexual desire leads you to copulate. Copulation leads to children. That's all the genes need. In the modern world contraceptives have been invented, so it's possible to enjoy copulation without the follow-up, without having children. And we do. And many of us do it all the time. And it is something which is manifestly and factually counter-Darwinian -- anti-Darwinian, anti the dictates of the selfish genes. We have been given brains which were shaped to enjoy sex. We have also been given brains that have been shaped to enjoy various other kinds of hedonistic pleasures. We have noticed consciously that hedonistic pleasure or other more worthwhile pastimes are sometimes incompatible with having lots of children all the time that you have to look after. And so we get the best of both worlds by consciously deciding to enjoy both the sex and the other things that would have been competed with -- by the need to look after children. We have achieved the best of both worlds from our own brain point of view, but not of course from the gene point of view.

QUESTION: Might religion itself be viewed as something that enhances survival?

MR. DAWKINS: I should have mentioned that religion may very well have a conventional Darwinian survival value. I have talked about religion as a virus, which is survival value at the level of the meme rather than the gene, the meme being the intercultural inheritance. But it could of course have survival value at the conventional gene level. Religion could enhance the survival of individuals who are religious. Or some people believe in some kind of group selection -- it could enhance the survival of a group. It's not difficult to see the advantages for a group, a tribe, which has a war-like god which enjoins its young men to sacrifice themselves for the good of the tribe, to fight with suicidal ferocity and bravery -- certainly it worked in the Crusades. Certainly it worked with the Japanese Kamikaze fighters. Certainly it works with Islamic suicide bombers. Such a tribe, such a group, could have great success in war, and therefore spread itself, spread the genes of the group, and of course spread the memes of the group, spread the ideas of the group.

But what about individual survival? It's possible too that religion might enhance individual survival. There is even some medical evidence that religious people in our society are slightly healthier on average than non-religious people -- perhaps because they suffer less from stress-related diseases. It's known that stress gives rise to disease. It's also known that many diseases, especially stress-related diseases, can be cured by placebos -- pills that have no medicinal effect, but people think they do, and so they do.

I could easily believe that religion could enhance health and hence survival, and that therefore there could be indeed be literally Darwinian survival value, Darwinian selection in favor of religion. None of that of course bears at all upon the truth value of the claims made by religions.

QUESTION: If you do not explain free will in a religious sense, how do you explain the ontology, the coming into being of free will?

MR. DAWKINS: The question of free will is a profound philosophical problem. And, as I said before, nothing is changed by adding the word "genes." So you could ask a question about how do you explain free will in a non-religious way -- not just to a Darwinian or somebody who believes that we are gene machines, but to anybody who takes a philosophically deterministic position.

So forget all about genes and think entirely about the philosophy of determinism. Now, everybody who calls themselves a determinist knows subjectively that they have the sensation of free will. We all know what it feels like to feel free. The only argument is whether fundamentally we are determined. That's one position which I wouldn't mind taking up. I don't find any difficulty with that position. I am quite prepared to believe that when I think I've taken the decision -- when I feel that I, with my own free will, have exercised a free choice, I've decided to do one thing rather than another -- I've decided to immigrate or decided not to immigrate, or to buy this house rather than that house -- it feels like free will. But it's perfectly possible that actually my decision to immigrate or not to immigrate was influenced by events in the brains which were influenced by other events, influenced by other events, which fundamentally all have a definite physical cause.

But it doesn't matter. There is no difference between the way it feels to have free will if there is this kind of fundamentally illusory free will that I've been talking about, or if in some other sense (which actually I can't quite imagine what it would like) we really did have free will. It wouldn't feel any different. It certainly wouldn't affect any of the arguments we ever have in selfish genery, when we ask whether we are free to depart from the dictates of the selfish genes and practice some anti-Darwinian behavior, such as contraception. Clearly that's no more difficult to imagine than deciding whether to immigrate or not to immigrate. The facts are that we do make such decisions. The facts are that we do take a decision to use contraception. And the -- contraception the mere existence of contraception -- something as mundane as that -- or the mere existence of the fact that one could take a decision to immigrate -- is not going to solve one of the great philosophical problems of all time. That's going to be there in any case. It doesn't actually affect how we feel or what we do.


Transposable-Element Vectors and Other Methods to Genetically Modify Drosophila and Other Insects

8.10 Evolution of Resistance to Transposable Elements

The spread of Ps into populations of D. melanogaster occurred worldwide since the 1930s. This invasion was remarkable because intact autonomous P elements induce severe disadvantages in the newly invaded populations. If P elements invade a small population, that population usually is lost due to the high rate of lethal genetic changes caused ( Engels, 1997 ). If evolution of repression systems (resistance to transposition) fails to occur quickly, invaded populations go extinct ( Corish et al., 1996 ).

Several P repressor systems (resistance mechanisms) were identified, illustrating the evolutionary importance of maintaining genome integrity from invasive TEs. The repressors are transmitted cytoplasmically (maternally inherited) or through the nuclear genome, in which case the transmission is biparental. The repressor systems have been classified as P, M’ or Q ( Corish et al., 1996 Badge and Brookfield, 1998 French et al., 1999 ). P fly strains have a strong maternally inherited repression system called P cytotype ( Engels, 1989 ). P cytotype is mediated by a 66-kDa protein produced by differential splicing of the complete element’s transcript ( Laski et al., 1986 ). When P females are crossed to a strong P line less than 10% of the ovaries are dysgenic, indicating P strains strongly repress hybrid dysgenesis. If P males are crossed to M females (which lack a repression system), more than 80% of the ovaries are dysgenic in their progeny. P strains are strong inducers of transposition. M’ strains also contain repressor elements. Repression in M’ strains is due to the KP element ( French et al., 1999 ). M’ strain females display intermediate levels of repression of dysgenesis when crossed to P males. Q strains strongly repress transposition. Some Q strains show a maternal mode of inheritance of repression while others have biparental mode. It thought a repressor (SR) results from a 308-bp deletion at the 3′ end of the P.

Resistance to P elements can develop rapidly, as demonstrated by two surveys of D. melanogaster populations along a 2800-km cline along the eastern coast of Australia. The first survey occurred in 1983 and the second in 1993. In 1983, P cytotype populations were found in the north, Q populations at central locations, and M’ populations in the south ( French et al., 1999 ). After 10 years, Q and M’ populations had increased their range at the expense of P lines.

In species of Drosophila in which P elements have been present a long time, no complete functional P has been found ( French et al., 1999 ). Instead, many populations contain tandem repeats of elements with degenerate fourth exons, which might encode repressor activity. In D. nebulosa, a complete element was isolated but the element contained many base changes in all four exons and was nonfunctional. These results reinforce the notion that active transposition of P is detrimental.

TEs consist of about 12% of the genome of D. melanogaster and are responsible for about 80% of spontaneous mutations ( Guerreiro, 2011 ). However, activity of TEs is not related to their abundance in the genome. Research to understand the triggers of transposition has focused on biotic and abiotic stresses and environmental changes. In D. melanogaster, a number of experiments suggest temperature and thermal stresses increase transposition rates, at least some of the time. Dysgenic and hybrid crosses, as well as colonization events, can increase transposition rates of some TEs. X-irradiation can initiate P-element transposition, and infections by viruses or other pathogens may, as well. The mutagen ethyl methanesulfonate (EMS) can activate P elements. Gamma irradiation appears to induce transposition of the 412 retrotransposon, but not of hobo, in D. melanogaster.